Medusae of the world

MEDUSAE OF THE WORLD

VOLUME I

THE HYDROMEDUSAE

BY

ALFRED GOLDSBOROUGH MAYER

WASHINGTON, D. C.

PUBLISHED BY THE CARNEGIE INSTITUTION OF WASHINGTON

1910

CARNEGIE INSTITUTION OF WASHINGTON
PUBLICATION No. 109, VOL. I

Copies of this Book
were first issuec*

AUG9 1910

CONTENTS.

VOLUME I.

Introduction i~4

Synopsis of Genera of the Hydromedusas 5-16

Medusae Milleporinae 16

Anthomedusae 17-196

Genera of the Anthomedusae:

Pachycordyle 21 Zancleopsis 91 Podocoryne 135

Amaithasa 22 Pteronema 92 Turritopsis 143

Pennaria 23 Eleutheria 93 Oceania 146

Trichorhiza 28 Mnestra 96 Stylactis 149

Steenstrupia 29 Ctenaria 98 Thamnostylus 151

Hybocodon 37 Cladonema 98 Thamnitis 152

Microcampana 44 Dendronema 102 Lymnorea 153

Dicodonium 44 Protiara 105 Bougainvillia 155

Sarsia 47 Heterotiara 107 Nemopsis 172

Stnuridiosarsia 64 Stomotoca 108 Rathkea • • '75

llydrichthys 66 Dissonema 115 Chiarella 182

Eucodonium 68 Pandea 116 Bythotiara ..185

Ectopltura 68 Turns I2O Sibogita 186, vol. 2 4QI

Corynitis 71 Conis 130 Niobia 187

Slabberia 73 Calycopsis 130, vol. 2 491 Proboscidactyla. . . . . . 188

Margelopsis 80 Cytsis ... ,:. .-...- ... 132 Willsia . . 193

Zanclea 85

Leptomedusae '..'.' • .196-230

Genera of the Leptomedusae:

Thaumantias 198. Staurodiscus 213 Dichotomia m

Laodicea 201 1'tvchogena 214 Dipleurosoma 224

Melicertum 207 I'olyorchis 218 Toxorchis 228

Melicertissa 209 Spirocodon 219 Netocertoides. . . .229

Orchistoma 21 r Cannota 221 Monobrachium. . --230

Timoides . . ..212 Cuvieria 221

MEDUSAE OF THE WORLD.

THE HYDROMEDUSAE.

INTRODUCTION.

This work was commenced in 1892 at the suggestion of Dr. Alexander Agassiz
while I was a student in his marine laboratory at Newport. Dr. Agassiz's plan
was that we produce conjointly a work upon the Medusae, Siphonophora, and
Ctenophorae of the Atlantic coast of North America. In pursuance of this plan,
Dr. Agassiz sent me upon expeditions to Halifax, Nova Scotia; Eastport, Maine;
Charleston, South Carolina; and Tortugas, Florida. It was also my privilege to
accompany him as his assistant upon his expeditions to the Bahamas, and to the
coral regions of the tropical Pacific. During these years the majority of our
American species were captured and drawings of them made from life.

The description of all of the then known American Atlantic forms was com-
pleted by me in 1900, but unfortunately the pressure of other and more important
work prevented the revision of the manuscript by Dr. Agassiz, and thus it remained
in the Museum of Comparative Zoology at Harvard University until 1904, when
Dr. Agassiz generously returned it to me with permission to publish it in any
manner whatsoever. Thus the original plan was reluctantly abandoned.

During the four years that elapsed while the manuscript lay unstudied at
Harvard, new forms had been discovered along our coast; and Nutting and Hargitt
had published their papers upon the hydroids and medusae of the Woods Hole
region.

It was necessary to thoroughly revise the manuscript, and in order to render it
of greater service, I have attempted to extend the original work to include descrip-
tions of all known forms of medusae from all parts of the world. This extension
was made possible through the generous establishment by the Carnegie Institution
of Washington of a Marine Research Laboratory at Tortugas, Florida. Many
forms were collected upon a cruise of the Carnegie Institution yacht Physalia
from Boothbay Harbor, Maine, to Tortugas, Florida; and new or interesting
medusae have been obtained each season upon excursions over the tropical Gulf
Stream, and among the Bahamas. My official position in connection with the
laboratory has afforded me every possible facility in time and opportunity tor tilt-
prosecution of these studies, and I can not too kindly express my sense of gratitude
to the executive officers of the Carnegie Institution tor their generous support.

To Geheimrath Prof. Dr. Anton Dohrn, and to his able corps of associates at
the Stazione Zoologica, I am indebted for numerous kindnesses shown to me during
my visit to the laboratory at Naples from November to February, 1907 and 1908.

I have also enjoyed full privileges of study in the libraries of the Museum of
Comparative Zoology at Harvard University; in the Boston Society of Natural

1

2 MEDUSvE OF THE WORLD.

History; the American Museum of Natural History; Columbia University, the
National Museum at Washington, and the Museum of the Brooklyn Institute of Arts
and Sciences. Through these facilities I have been enabled to review nearly all
of the published works upon medusae, but the review of literature can not pre-
tend to completeness for 1907 and 1908, although all papers of those years which
the author could discover are recorded.

Moreover, Profs. William K. Brooks and Louis Murbach have been so kind
as to lend some of their original drawings, which are reproduced in this work, and
the following gentlemen have generously granted permission for the reproduction
of figures from their published works, thus enabling us to present text-figures of
many forms which would otherwise have been represented merely by descriptions:
Dr. Alexander Agassiz, Director of the Museum of Comparative Zoology at Harvard
University; Dr. Henry B. Bigelow, of Harvard; Dr. R. P. Bigelow, of the Massa-
chusetts Institute of Technology; Prof. Edward T. Browne, of the University of
London; Prof. Dr. Carl Chun, of Leipzig; Prof. Dr. S. Goto of Tokyo;
Geheimrath Prof. Dr. Ernst Haeckel, of Jena; Prof. C. W. Hargitt, of Syracuse
University; Prof. Dr. Cl. Hartlaub, of Helgoland; Prof. W. C. M'Intosh, of Aber-
deen University; Prof. Dr. Otto Maas, of Miinchen; Prof. C. C. Nutting, of Iowa;
Prof. Henry F. Perkins, of Vermont; and Prof. Dr. Ernst Vanhoffen, of Kiel.

I have always felt that each working naturalist owes it as a duty to science to
produce some general systematic work, and this has been an actuating motive
in the production of this book. But chiefly have I been moved to the task through
respect for the wishes of my generous friend and master in science, Alexander
Agassiz. Nor can one remain insensible to the rare grace of form and delicate
beauty of color of these creatures of the sea, associated as their study is with mem-
orials of the labors of a host of distinguished naturalists. Dry though these pages
must be to the reader, to the writer they are replete with memories of the ocean
in many moods, of the palm-edged lagoons of coral islands sparkling in the tropic
sun, of the cold, gray waters of the northern sea bestrewn with floating ice, of days
of withering calm in the heat of the torrid zone, and of adventure in the hurricane ;
all centering around the absorbing study of the medusa?. Love, not logic, impels
the naturalist to his work.

This book attempts to present a new classification of the medusae. With every
respect for Haeckel's great work, it has appeared to me that its subdivisions are
often too precise to be convenient, and too artificial to accord with nature. More-
over, many of Haeckel's genera are founded upon intergrading characters, and are
thus imperfectly separated. The young of many medusae appear in one genus,
and the adults in another. The aim of Haeckel's system is to emphasize distinctions,
whereas my aim is to indicate relationships. I therefore attempt to separate genera
upon positive, not upon relative, distinctions. For example, if we define one genus
as having a narrow manubrium (Margelis), and another as having a wide manu-
brium (Boiigamvillta), we must either institute a third genus for newly-discovered
medusae with manubria of moderate width or place them doubtfully in one or the
other of the genera of the extreme members of the series.

On the other hand, if we define one genus as having eight tentacles, and
another as having nine or more tentacles, there can be no confusion between them,
for the difference, although slight, is positive and numerical, not qualitative and
intergrading.

I have not described hydroids which do not produce free-swimming medusae,
although I grant it is wholly illogical to admit Podocoryne into a system which
excludes Hydractmia, or to include only those species of Stylactis which produce

INTRODUCTION.

medusae and to exclude those which produce sessile gonophores. Nevertheless
this should be clearly understood and must be accepted as an artificial limitation
of the work.

I have thus attempted to describe only such hydroids as are known to produce
medusae, and have endeavored to bring the systematic arrangement of the medusa
more nearly into accord with that of the hydroids. A strictly natural system includ-
ing both hydroids and medusae can not be constructed, for many of the hydroids
remain undetermined. Moreover, dissimilar hydroids (Svm-tjrv>if, StanriJinim
may give rise to similar medusae (Sarsia); or the reverse may be the case, as in the
medusae of Bongamvillia and Nemopsis, or that of the two sorts of medusa- (Sarsia
and Cladoncrna] which may arise from hydroids of Stauridia.

These and many other cases of a similar nature interpose a barrier to our
attempts to invent a natural system which includes all hydroids and medusa" within
its embrace. At present, I believe, we must content ourselves with a compromise
between a natural and an artificial arrangement, confiding in the belief that as
more and more of the hydroids are discovered it will become correspondingly
more possible to arrange the medusae in a natural system. After consultation with
Prof. C. C. Nutting we have mutually decided that the promulgation of such ;i
system is at present inadvisable. Such a system has, indeed, been proposed by
von Lendenfeld, 1884 (Zool. Anzeiger, Bd. 7), but has gained no acceptance.

Much confusion has been introduced through the habit, in vogue among
marine expeditions, of sending all of the medusa" to one specialist and the hydroids
to another. Thus the sessile and the reproductive stages of the same animals are
worked upon independently from different view-points by different men.

I am inclined to regard the Trachymedusae and Narcomedusa" as being
transformed actinutae, for they commonly develop through an actinula larva in
which the bell grows out as a collar-like, or intertentacular lappeted expansion
from the sides of the body after the tentacles have appeared, and the tentacles of
the actinula become those of the medusa. The medusa of the Anthomedusas and
Leptomedusae is formed upon a different plan, for the tentacles grow outward from
the bell-margin after the bell has developed. I believe, therefore, that the bell of
the Trachymedusae and Narcomedusae is not homologous with that of the Antho-
medusa and Leptomedusae. It is evident that the entodermal otoliths of the
Trachymedusa" and Narcomedusae are not homologous with the ectodermal otoliths
of Leptomedusae. Budding and alternation of generations occur in both classes
of veiled medusae.

I believe that the medusa-shape has been acquired independently in the
Trachyhna and Leptolma forms of veiled medusae.

The colored plates contained in this volume consist of drawings, from life, of
medusae of the Atlantic coast of the United States. The text-figures, on the other
hand, are chiefly outline tracings from the illustrations of many authors; and are
presented in order to spare the reader the trouble of consulting numerous scattered
works of reference. These outline copies of previously-published drawings of
medusae were carefully traced from the originals by Mr. Carl Kellner, artist of the
Tortugas Marine Laboratory of the Carnegie Institution.

Other outline figures are from life, and the majority of these were drawn by
the author while studying at Mousehole, Cornwall, England, and at the Naples
Zoological Station during the autumn and winter of 1907 and 1908.

This book aims to be something more than an old-fashioned systematic treatise,
for it attempts to record, if not to review, all works upon the embryology, cytology,
oecology, physiology, etc., of all forms coming within the scope of the text.

MEDUSAE OF THE WORLD.

Many of the medusae are modified more or less profoundly by environmental
conditions, and this gives rise to numerous local races, the determination of the
relationships between which is all but impossible in the present state of our knowl-
edge. Experimental work in this field is in its infancy, yet the few observations of
Hallez on Bongainvillia and of Browne in his experiments in maintaining various
hydroids under different conditions in aquaria suggest that the results of more
extended studies will be of great benefit to the systematist in determining the
natural limitations of species. A striking example of the profound effect of environ-
mental changes is seen in the experiments of Goldfarb, 1906, upon Eudendrium,
wherein he showed that after the regenerative process had entirely ceased in the
dark, it could be recalled into activity by an exposure of only five seconds to the
daylight.

At present the natural classification of the species of such genera as Obclia,
Phialidnnn, Etttnna, Bougamvillia, etc., is impossible, and the Linnean system is
inadequate to the task of expressing their actual relationships. Indeed, with the
exception of the sponges and corals, there is no phylum of the animal kingdom more
difficult to classify than the medusae.

In a work of this magnitude there must needs be both errors and omissions,
and for these I can but present my apologies in advance of their discovery, trusting
that all such will soon be discovered and announced, and that science may be more
advanced than hindered through the publication of this book. I trust that none
of my esteemed contemporaries will feel aggrieved at such criticisms of their labors
as may appear in the following pages, for should anyone find cause for such offense,
he may console himself in the fact that I am obliged to confess to having found
more to criticize and amend in my own previously-published papers than in those
of any other living naturalist.

ANTHOMEDUS^:.

HYDROMEDUS/E OR VEILED MEDUS/E.

Medusae with a velum, or diaphragm, which partially closes the marginal opening of
the bell. When ripe the sexual products are found in the ectoderm.

With a double marginal nerve-ring, one above and one below the velum.

Without gastric filaments. Development either direct from actinula larvae or through
alternation of generations from hydroids.

Order 1. ANTHOMEDUS/E Haeckel, 1879.

Hydromedusae with gonads in the ectoderm of the manubrium. Without otocysts.
The hydroids are of the Tubularian order.

Family No. i, CODONID.&.

Anthomedusae with ring-like gonad encircling the manubrium. Four to six simple,
unbranched radial-canals. Simple, unbranched tentacles.

Subfamily No. i, SARSIANJE.
Some or all of the tentacles arise singly, not in clusters, from bell-margin.

PACHYCORDYLE, Weismann, i8%$ = Parvanernus ( ?), Mayer, I9°4-

Degenerate medusae without tentacles, radial or circular canals. Hydroid: Pachy-
cordyle Weismann.

AMALTH^EA, Schmidt, 1854.

Four rudimentary tentacles, four radial-canals, and a ring-canal. Hydroid: Corymorpha.

PENNARIA, Oken, 1815; Goldfuss, i%2O = Globiceps, Haeckel, 1879.
Similar to Amalthaa, but the hydroid is Pennana.

TRICHORHIZA, Russell, 1906.

Medusa resembles Pennaria, but the hydroid is Tnchorhiza.

STEENSTRUPIA, Forbes, iS^.6 = Eup/i\s(i, Forbes, i$4.% = Eupliysora, Maas, 1905.

Four radially situated tentacles, one of which is long, the other three short. Bell radially

symmetrical. Hydroid: Corymorpha.
HYBOCODON, L. Agassiz, \%()2 = HybocodonJt-Amphicodon, Haeckel, 1879.

One or more well-developed tentacles arise from base of one of the four radial-canals.
With rudimentary tentacles at the bases of the three other radial-canals. Bell asym-
metrical. Hydroid: H ybocodon Agassiz.

MICROCAMPANA, Fewkes, 1889.

With one long and five short tentacles, arising at ends of six radial-canals.

DlcoDONlUM = Dicodonium + Dincmti, Haeckel, 1879.

Two well-developed and two rudimentary tentacles. No meridional lines of nettle-cells

over exumbrella.
SARSIA, Lesson, i^^.^ = CoJontutn + Sarsia + S'vnJii'tyon, Haeckel, 1879.

Four equally developed tentacles with abaxial, ectodermal ocelli. No meridional lines

of nettle-cells over the exumbrella. Hydroid: Syncoryne.
StauriJiosarsia new subgenus. Medusa similar to Sarsia, but the hydroid is Stuundia.

HYDRICHTHYS, Fewkes, 1888.

Medusa resembles Sarsia, but there are no ectodermal ocelli upon the tentacle-bulbs.
Hydroid: H vJrichthys, Fewkes; commensal or parasitic upon fish.

EUCODONIUM, Hartlaub, 1907.

Medusa resembles Sarsia, but stomach is mounted upon a gelatinous peduncle. The
four tentacles terminate each in a knob-shaped extremity.

ECTOPI.EURA, L. Agassiz, 1862 (sens. ampl.).

With two or four tentacles. Eight meridional lines of nettle-cells over the exumbrella.
Hydroid: Ectopleura.

6 MEDUS.E OF THE WORLD.

CORYNITIS, McCrady, 1857.

Four knobbed tentacles. Manubrium cruciform in cross-section. Ocelli upon tentacle-
bulbs. Hydroid: Syncoryne (Hargitt).

SLABBERIA, Forbes, lB^6 = Slabberia + Dif>urena + BathycoJon, Haeckel, 1879.

Four knobbed tentacles. Manubrium tubular and encircled by two or more ring-like
gonads. Hydroid: Syncoryne.

Subfamily No. 2, MARGELOPSIN^E.

Four radially placed clusters ot marginal tentacles. No oral tentacles. A ring-like
gonad encircles the stomach. Four simple, unbranched radial-canals. Hydroids are pelagic
Tubularians, and the medusae arise by budding from their sides.

MARGELOPSIS, Hartlaub, 1897.

Characters of the medusa are those of the subfamily. Hydroid: Margelopsis, in which
the tentacles are disposed in definite circlets.

PELAGOHYDRA, Dendy, 1903.

Medusa similar to Margelopsis. Hydroid: PelagohyJra. Its tentacles arise irregularly
from sides of the hydranth, and are not disposed in definite circlets.

Family No. 2, CLADONEMID^E.

Tentacles branch dichotomously or complexly, or give rise to a linear series of nematocyst-
bearing filaments along their abaxial sides. Gonads ring-like, or segregated upon the inter-
radial and adradial sides of the manubrium.

Subfamily No. i, PTERONEMINJE.

Manubrium without oral tentacles.

ZANCLEA, Gegenbaur, l8^6 = Zanclea + Gernmaria, Haeckel, 1879.

With two or four tentacles, each of which gives rise to an abaxial row of nematocyst-
bearing branches. With meridional rows of nettle-cells upon the exumbrella.
No ocelli upon the tentacle-bulbs. No brood-pouch above the stomach. Hydroid:
Gemmaria McCrady; Allman.

ZANCLEOPSIS, Hartlaub, 1907.

Similar to Znticlea, but without meridional lines of nettle-cells over the exumbrella. With
ectodermal ocelli upon the outer sides of the tentacle-bulbs.

PTERONEMA, Haeckel, 1879.

Similar to Zanclea, but with a brood-sac above the stomach. Four tentacles.

ELEUTHERIA, Ouatrefages, 1842.

Four to six bifurcated tentacles, and an equal number of simple unbranched radial-
canals. There is an ectodermal brood-sac above, but not connected with the stomach.
The medusa is hermaphroditic and the germ-cells develop in the brood-sac. Hydroid:
Clavatella Hincks.

MNESTRA, Krohn, 1853; Giinther, 1903.

Degenerate medusa parasitic upon Phylhrrhoe. Four to no tentacles, each with an
abaxial line of nettle-warts. Four perradial meridional lines of nettle-cells over
the exumbrella. Four radial-canals. Throat is blocked by a cavernated mass of
entoderm.

Subfamily No. 2, DENDRONEMIN^).

Manubrium with oral tentacles.

CTENARIA, Haeckel, 1879.

Two marginal tentacles with abaxial filaments. Four bifurcated radial-canals. Simple
oral tentacles. Brood-sac above the stomach.

ANTHOMEDUS.E. 7

CLADONEMA, Dujardin, 1843.

Four or five bifurcated or eight to ten simple radial-canals. Branched marginal tentacles.
Simple oral tentacles. No brood-sac above the stomach. Hydroid: Statin Jin
Dujardin.

DENDRONEMA, Haeckel, 1879.

Similar to ClaJonema, but with branched oral tentacles and with brood-sac above stomach.

Family No. 3, OCEANID.S, sensu Vanhoffen.

Anthomedusae in which the gonads are segregated and developed upon the interradial
or adradial sides of the manubrium. With unbranched marginal tentacles. Mouth with
four lips.

Subfamily No. i, TIARIN.E.

Unbranched radial-canals. Marginal tentacles separate; not grouped into clusters. No
oral tentacles. Tentacles hollow. When present the ectodermal ocelli are upon the abaxial
sides of the tentacle-bulbs. With the exception of Calycopsis all of the genera have four
radial-canals.

PROTIARA, Haeckel, i%jq = Halitiara, Fewkes, 1882.

Four radially placed, well-developed tentacles. Four interradial gonads with smooth
outer surfaces. With or without marginal cirri. External surfaces of gonads smooth.
Four cruciform, simple lips. No ocelli on the velar sides of the tentacles.

HETEROTIARA, Maas, 1905.

Eight or more tentacles. The ring-canal gives rise to blindly-ending centripetal vessels.

STOMOTOCA, L. Agassiz, \%()2 = Amf>hinema-\-Stomotoca + Codonorchis, Haeckel, 1879.

Two well-developed and many rudimentary tentacles. External surfaces of the adradial
gonads are thrown into transverse folds. Hydroid: Pengonimus.

DISSONEMA, Haeckel, 1879.

Similar to Stomotoca, but the gonads finally migrate outward along the four radial-canals.

PANDEA, Lesson, 1843.

Four or more tentacles. Gonads four interradial, folded ridges on the sides of the
stomach but these gonads are not completely separated in the four principal radii.
Hydroid: Dcndroclava ( ? ?).

TURRIS, Lesson, 184.3 = Tiara + Turris + Catablema, Haeckel, 1879.

Four or more tentacles. Four interradial horseshoe-shaped gonads on the stomach
wall. These are composed of more or less fused ridges or network-like swelling.
They are completely separated in the four principal radii. Hydroid: Clirculu Wright.

CONIS, Brandt, 1834; Haeckel, 1879.

Similar to Pandea, but the tentacle-bulbs give rise to abaxially-placed clubs which bear
ocelli.

CALYCOPSIS, Fewkes, 1882.

Sixteen simple, separate radial-canals. Eight transversely folded, adradial gonads.
Ring-canal simple.

Subfamily No. 2, MARGELINjE.

With four unbranched radial-canals. With oral tentacles, or nematocyst-knobs, upon
the lips. Tentacles solid. When present the ectodermal ocelli are upon the inner (velar)
sides of the tentacles.

CYTVEIS, Eschscholtz, iS2() = Cytxis + Cubogastcr, Haeckel, 1879.

Four simple marginal tentacles. With simple, unbranched, oral tentacles.

PODOCORYNE, Sars, \?>Jtb = Dysmorf>hosa-\-Cytteandra, Haeckel, 1879.

Eight or more simple marginal tentacles. With simple, unbranched, oral tentacles.
Hydroid: Podocoryne. When present the peduncle above the stomach is solid and
gelatinous.

MEDUSAE OF THE WORLD.

TURRITOPSIS, McCrady, 1856.

Eight or more simple marginal tentacles. The entodermal walls of the radial-canals
above the stomach are composed ot vacuolated cells forming a peduncle-like base
for the stomach. The mouth is studded with a row of nematocyst-bearing knobs.
With ectodermal ocelli on the velar sides of the tentacles near their bases. Hydroid:
DenJroclava (Brooks).

OCEANIA sensu Kolliker, 1853; Gegenbaur (in part) 1856.

Medusa similar to Turritopsis, but with solid gelatinous, non-vacuolated peduncle above
the stomach. Hydroid: C lava-like.

STYLACTIS, Allman, 1864.

Degenerate medusae, with four to eight rudimentary marginal tentacles and no oral
tentacles. Hydroid: Stylactis.

THAMNOSTYLUS, Haeckel, 1879.

With two simple, marginal tentacles, and with branched oral tentacles.

THAMNITIS, Haeckel, 1879.

Four radially placed, simple, marginal tentacles, and branched oral tentacles.

LYMNOREA, Peron and Lesueur, iSog — Limnorea + Thamnostoma, Haeckel, 1879.
Eight or more simple, marginal tentacles. Branched oral tentacles.

BOUGAINVILLIA, Lesson, \$4.T> = Margelis + Lizusa + Hif>pocrene, Haeckel, 1879.

With branched oral tentacles. The marginal tentacles are grouped in four radial clusters.
All of the tentacles are filiform. Hydroid: Bougainvillia.

NEMOPSIS, L. Agassiz, 1849.

Similar to Bougainvillia, but each cluster of marginal tentacles consists of a median
pair of clavate tentacles flanked by filiform tentacles. Hydroid: Bougainvillia.

RATHKEA, Brandt, lS^y = Lizzia + Lizella + Rathkea + Margellium, Haeckel, 1879.

With eight clusters of marginal tentacles. Simple or branched oral tentacles. Ring-
canal simple.

CHIARELLA, Maas, 1897.

Sixteen (eight double) clusters of marginal tentacles. The ring-canal gives rise to centrip-
etal vessels. Branched oral tentacles.

Subfamily No. 3, DENDROSTAURINjE.

With branched radial-canals. No oral tentacles. Marginal tentacles arise singly, and
are not grouped into clusters. Tentacles hollow. No cirri or marginal clubs.

BYTHOTIARA, Giinther, 1903.

Four bifurcated radial-canals and a ring-canal. Four interradial gonads.

SIBOGITA, Maas, 1904.

Four principal radial-canals, which branch complexly. Four interradial gonads. Ring-
canal present.

NIOBIA, Mayer, 1900.

Four principal radial-canals, two of which bifurcate so that six canals reach the circular
vessel. Four interradial gonads. The marginal tentacles develop into medusae,
and are cast off.

PROBOSCIDACTYLA, Brandt, \%Tfi=Dyscannota + Dicranocanna+ Willeta

-\-Proboscidactyla, Haeckel, 1879.

The four primary radial-canals give rise to simple or branched side branches. No ring-
canal. With intertentacular lines of nematocysts upon the exumbrella above the
bell-margin. Gonads on the adradial sides of the stomach extending outward
along the sides of the four main radial-canals.

LEPTOMEDUS.E.

9

WILLSIA, Forbes, 1846.

Similar to ProbosciJactyla, but with six or more primary radial-canals. The hydroid
belongs to the genus Lar Gosse.

Order 2. LEPTOMEDUSjE Haeckel, 1886.

Hydromedusae with gonads upon the radial-canals. When present the otoliths are of
ectodermal origin. The medusae arise through alternation of generations from Campanu-
larian hydroids.

Family No. i, THAUMANTIAD^).

Leptomedusae without lithocysts.

Subfamily No. i, MELICERTIN^.

With simple, unbranched radial-canals and an equal number of lips, without oral ten-
tacles. Cirri or marginal clubs may or may not be present.

THAUMANTIAS, Eschscholtz, \%29 = Tetrtinfmti + T/iiiii>tiiinti(is, Haeckel, 1879.

Four or more tentacles. Four radial-canals. No marginal clubs or cirri. Hydroid:
T haumantias.

LAODICEA, Lesson, i$4$ = Octonema + Laodice, Haeckel, 1879.

Four or more tentacles. Four radial-canals, with marginal clubs or cirri. Hydroid:
Cuspidella.

MELICERTUM, Oken, 1815; sensu L. Agassiz, i%ta=Melicertella+Melicertum, Haeckel, 1879.
With eight or more tentacles. Eight radial-canals. No marginal clubs or cirri. Hydroid:
Melicerturn Agassiz.

M.ELlCER.TlSSA = Mclicertissa + MflicfrtiJnitn, Haeckel, 1879.

Similar to Melicertum, but with marginal clubs or cirri between the tentacles.

ORCHISTOMA, Haeckel, 1879.

More than eight radial-canals. With or without marginal clubs or cirri.

TIMOIDES, H. B. Bigelow, 1904.

Four radial-canals. The ring-canal gives rise to blindly-ending centripetal diverticula.
Numerous tentacles and cirri. Four gonads on the four radial-canals. Stomach
mounted upon a peduncle.

Subfamily No. 2, POLYORCHIN.E.

The radial-canals give rise to branches which end blinJlv and do not connect with the
marginal ring-canal.

STAURODISCUS, Haeckel, 1879.

With four radial-canals, each of which gives rise to two side branches, which end blindly.

PTYCHOGENA, A. Agassiz, 1862, 1865.

With four radial-canals which give rise to numerous blindly-ending side branches. Gonads
leaf-shaped and developed upon the side branches of the radial-canals.

POLYORCHIS, A. Agassiz, 1862 to 1865.

With four radial-canals which give rise to numerous blindly-ending side branches.
Numerous sac-like, sausage-shaped gonads attached to the radial-canals and to
their side branches. Ring-canal simple. Bell-margin not cleft into lappets.

SPIROCODON, Haeckel, lS^g = Goniomeam/rus, Kirkpatrick, 1903.

Similar to Polyorchis, but the ring-canal gives rise to blindly-ending, centripetal branches,
and bell-margin is cleft into lappets.

10 MEDUS/E OF THE WORLD.

Subfamily No. 3, BERENICINJE.
The radial-canals give rise to branches which connect with the ring-canal.

CANNOTA, Haeckel, 1879.

With four radial-canals, each of which gives rise to two side branches which join the
ring-canal.

CuviERIA, Peron, 1807 = Berenice, Haeckel, 1879.

With four main radial-canals which give rise to numerous non-dichotomous side branches.
Gonads on the terminal ramuli of the canals.

DICHOTOMIA, Brooks, 1903.

With four main radial-canals which divide dichotomously two or more times. The
gonads extend outward from the stomach over the canals.

DIPLEUROSOMA, Axel Boeck, i$66 = Tetracannota, Mayer, 1900.

With three or more main radial-canals which give rise to non-dichotomous side branches.
Gonads on the canals adjacent to the stomach. Hydroid: Cuspidella (?).

ToxoRCHiS = Toxorchis + ClaJocanna, Haeckel, 1879.

Four or more main radial-canals which branch dichotomously one or more times. Gonads
on the outer branches of the canals near the ring-canal.

NETOCERTOIDES, Mayer, 1900.

Eight main radial-canals which branch dichotomously. Gonads extend outward from
the sides of the stomach along the radial-canals.

Family No. 2, EUCOPID^.

Leptomedusaewith lithocysts, and with less than eight radial-canals upon which the gonads
are developed.

Subfamily No. i, OBELIN^E.
With eight adradial lithocysts. Four radial-canals. Stomach without a peduncle.

EUCOPELLA, von Lendenfeld, 1883.

Degenerate medusae. No tentacles. No manubrium. Branched radial-canals. Hydroid:

Eucopella.

AGASTRA, Hartlaub, 1897.

Degenerate medusae. No manubrium. Simple, unbranched radial-canals. Hydroid:
Campanularia ( ?).

EUCOPE, Gegenbaur, 1856.

Basal bulbs of the tentacles are simple and hollow, and do not project inward into the
gelatinous substance of bell. Lithocysts on bell-margin. Hydroid: Campanulana.

OBELIA, Peron and Lesueur, 1809.

Entodermal cores of tentacles project inward into the gelatinous substance of the bell.
Otocysts on bases of tentacles. Hydroid : Obelia.

TIAROPSIS, L. Agassiz, 1849.

An ocellus with entodermal pigment above each lithocyst. Tentacle-bulbs simple and
hollow.

Subfamily No. 2, PHIALIN^.

With more or less than eight lithocysts. Four to five radial-canals. Stomach without a
peduncle.

CLYTIA, Lamouroux, \%i2 = Ef>enthesis, McCrady, 1857.

Sixteen tentacles alternating with sixteen lithocysts. Four radial-canals. No cirri.
Hydroid: Clytia.

LEPTOMEDUS.E. 11

PHIALIDIUM, Leuckart, i%$() = 0<:eania, Agassiz, 1862, 1865.

Sixteen or more tentacles. More than sixteen lithocysts. Four radial-canals. No rudi-
mentary tentacles. No cirri. Hydroid: Campanulina.

PHIALUCIUM, Maas, 1905.

Similar to Phialidium, but with permanently rudimentary tentacle-bulbs. Hydroid un-
known.

BLACKFORDIA, Mayer.

Numerous tentacles and lithocysts. Entodermal cores of some or all of the tentacles
project inward into the gelatinous substance. Four radial-canals. No cirri. No
permanently rudimentary tentacles.

PSEUDOCLYTIA, Mayer, 1900.

Five radial-canals, five lips, five gonads. Numerous tentacles and lithocysts. No cirri.

GASTROBLASTA, Keller, \?>?>^=Multioralis, Mayer, 1900.

Two or more manubria. No cirri. The medusa propagates by fission.

EUCHEILOTA, McCrady, 1857.

Four or more closed vesicular lithocysts. Four radial-canals. Marginal or lateral cirri.
Hydroid: Campanulina ( ?).

MnROCOMA = Phialis + Mitrocoma + Mitrocomiurn + Mitrocomella, Haeckel, 1879.

Similar to Eucheilota, but the lithocysts are contained in open folds of the velum. No
entodermal ocelli such as are found in Tiaropsis.

STAUROPHORA, Brandt, lS^ = Staurostoma + Staurof>hora, Haeckel, 1879.

Mouth an open, cruciform, gutter-like slit extending down the four radial-canals.

Subfamily No. 3, EUTIMINJE.

With eight adradial lithocysts. Stomach mounted upon a gelatinous peduncle. Four
radial-canals.

SAPHENIA, Eschscholtz, 1829.

Two tentacles. Numerous cirri.

EUTIMA, McCrady, 1 857 = Eutima + Eutimeta + Octorchis + Octorchandra

-\-Eutnnalphcs, Haeckel, 1879.

Four or more tentacles. Numerous cirri or marginal warts. Four or eight gonads upon
the four radial-canals. Hydroid: Campanopsis (Claus, Brooks).

JLwTIMlUM = Eutimium + Octorchi(lium, Haeckel, 1879.

Similar to Eutima, but without cirri. Hydroid: Campanulina ( ?)

Subfamily No. 4, EIRENINvE.

With more than eight lithocysts. Stomach mounted upon a gelatinous peduncle. Four
or six radial-canals.

PHORTIS, McCrady, 1857.

Four or more tentacles. No cirri. Four radial-canals. Hydroid: P/iortis Brooks.

IRENOPSIS, Goette, 1886, non Ireniopsis, Mayer, 1894.

Six or more tentacles. Six radial-canals. Six lips. Six gonads.

EIRENE, Eschscholtz, i%2g= Irene +Irenittm, Haeckel, 1879.

Four or more tentacles. Numerous marginal warts or cirri. Four or eight gonads
developed upon limited parts only of the four radial-canals. Hydroid: Campan-
ulina ( ?)

TIMA, Eschscholtz, 1829.

Similar to Eirene, but with gonads upon the entire lengths of the four radial-canals.
Hydroid: Campanulina ( ?)

12 MEDUS.E OF THE WORLD.

Family No. 3, ^EQUORIDjE.

Leptomedusae with otocysts, and with eight or more radial-canals.

OCTOCANNA, Haeckel, 1879.

Eight radial-canals, 45° apart. Eight lips. No ocelli. (Is this a young jEquorea ?)

OCTOGONADE, Zoja, 1896.

Similar to Octocanna, but the marginal sense-organs have ocelli as well as lithocysts.

STOMOBRACHIUM, Brandt, lS^^ = Stomobrachium + Staurobrac/iium, Haeckel, 1879.

Eight or more simple, unbranched radial-canals, which arise at equal intervals from the
margin of the stomach. Four lips.

HALOPSIS, A. Agassiz, 1863, 1865.

Radial-canals arise in four groups from the four perradial corners of stomach. Four lips.

^EQUOREA, Peron and Lesueur, i$og = &(juorea + R/iegmatoJes + Mesonema

-\-Polycanna, Haeckel, 1879.

More than eight simple, unbranched radial-canals which arise separately from the mar-
gin of the stomach. More than four lips. Subumbrella smooth, without gelatinous
papilla-like protuberances. Hydroid: Campanulina.

ZYGODACTYLA, Brandt, 1835; sensu Agassiz, 1862.

Similar to jEquorea, but with interradial rows of papilla-like, gelatinous protuberances
upon the subumbrella.

, Haeckel, 1879.

With bifurcated or branched radial-canals which arise at equal intervals from the margin
of the stomach. More than four lips. No peduncle.

ZYGOCANNULA, Haeckel, 1879.

Similar to Zygocanna, but the stomach is mounted upon a gelatinous peduncle.

Order No. 3. TRACHYMEDUS.E Haeckel, 1866.

Medusae with a marginal velum, and with lithocyst concretions of entodermal origin.
With simple uncleft bell-margin.

Family No. i, OLINDIADJE.

Some or all of the tentacles project from the sides of the bell, above the margin, and have
adhesive disks. Gonads linear, sac-like, or folded, and developed upon the four or six radial-
canals. The tentacles arise separately and are not grouped in clusters.

GONIONEMUS, A. Agassiz, 1862, 1865.

All of the tentacles project from sides of bell in a zone slightly above bell-margin. All
have adhesive disks. Four radial-canals. Lithocysts external, on bell-margin. No
centripetal canals. Development through an attached hydra stage. (Perkins.)

CUBAIA, Mayer, 1894.

Similar to Gonionemus, but with two sets of tentacles, one arising from the bell-margin
and the other set projecting from the sides of the bell, as in Gonionemus.

VALLENTINIA, Browne, 1902.

Similar to Cubaia, but with lithocysts inclosed within the gelatinous substance of the
bell, adjacent to the ring-canal, and on the inner side above the velum. (Is this
a young Olindias])

OLINDIAS, F. Miiller, 1861.

Similar to / allentmia, but with blindly-ending, centripetal diverticula from the ring-canal.

TRACHYMEDUS.K. 13

OLINDIOIDES, Goto, 1903.

Similar to OlinJias, but with six radial-canals (two bifurcated and two simple). Six
gonads. Four lips. The exumbrella tentacles project at various levels from the
sides of the bell.

Family No. 2, PETASIDjE Haeckel, 1879.

Trachymedusae with tour radial-canals upon which the linear or sac-like gonads are
developed. Tentacles without adhesive disks. Four lips.

PET ASUS = Pftasus + Dif>etasus + Petas(ita + Pftacli>: uni, Haeckel, 1879.

Tentacles arise at equal intervals, not grouped into clusters. No centripetal canals.
Free marginal hthocyst-clubs.

AGLAUROPSIS, F. Miiller, 1865.

Similar to Petasus, but the lithocysts are vesicular, and project from the bell-margin
between the tentacles.

CRASPEDACUSTA, Lankester, i$%o=LimnocoJium, Allman, 1880.

Tentacles arise singly as in Pi-tnsus and Aglauropsis, but the lithocyst concretions are
each inclosed in a cavity within the gelatinous substance of the velum on the inner
(centripetal) side of the ring-canal. (The medusa lives in fresh water among water-
lilies.) The hydroid is devoid of tentacles.

MlCROHYDRA, PottS, 1885.

Is possibly identical with Limnocodiutn, but the mature medusa is unknown. The
young medusa has no lithocysts, and it arises by budding from a minute hydroid
which has no tentacles.

MJ^OTIAS, OstroumofF, 1896.

Tentacles arise at equal intervals, not in clusters. Numerous centripetal, blindly-ending
canals arise from the ring-canal.

GOSSEA, L. Agassiz, 1862.

Tentacles grouped into clusters. No centripetal canals. Lithocyst concretions free or
inclosed.

Family No. 3, LIMNOCNIDIDjE.

Numerous hollow tentacles which project singly, not in clusters, from the sides of the
bell in a zone slightly above the margin. Tentacles without adhesive disks. Numerous
inclosed lithocysts on the exumbrella side of the velum. Mouth a round opening. Gonads
developed diffusely in the ectoderm of the stomach-wall. Four (occasionally five or six) radial-
canals. Medusa-buds arise from the sides of the stomach, and are set free.

LIMNOCNIDA, Giinther, 1893.

Generic characters are those of the family. The only known species is /,. tanganjicd
from the fresh-water lakes of Central Africa, and the Niger river.

Family No. 4, PTYCHOGASTRIDJE.

Numerous more or less isolated clusters of tentacles, some of which bear adhesive disks.
Numerous free lithocyst-clubs. Eight radial-canals. Four lips. Stomach eight-lobed. These
stomach-lobes are in the radii of the radial-canals, and are bound to the subumbrella by
means of eight mesenterial partitions. The gonads are upon the eight stomach-lobes, and
each is more or less divided by the mesentery so there may be eight double (sixteen) gonads.

PTYCHOGASTRIA, Allman, iSfi^Pcctyllis + Pectis + Pectanthis, Haeckel, 1879.
The generic characters are those of the family.

Family No. 5, TRACHYNEMID^E.

Trachymeduss with eight simple radial-canals upon which the gonads are developed.
No mesenterial partitions in the subumbrella. Tentacles without adhesive disks. Ring-
canal simple without centripetal branches.

14 MEDUSA OF THE WORLD.

Subfamily No. i, RHOPALONEMINjE.

Trachynemidae in which the stomach lacks a peduncle.
RHOPALONEMA, Gegenbaur, iS^6=Trachynema (young medusa), Gegenbaur, 1854

= Trachynema + Rhopalonema + Marmanema, Haeckel, 1879.

With eight well-developed radial tentacles, and eight or more small cirrus-like or club-
shaped interradial tentacles. All tentacles arise in a single row. Eight gonads
localized on the eight radial-canals. Four lips.
SMINTHEA, Gegenbaur, 1856.

Similar to Rhopalonema, but with only eight tentacles, one at the foot of each of the

eight radial-canals.

HOMCEONEMA, Maas, i8g3 = Colobonema, Vanhoffen, igo2 = lsonema (in part), Maas, 1906.
Similar to Rhopalonema, but the tentacles are all of one sort. No small club-shaped or

cirrus-like tentacles. Four lips.
PANTACHOGON, Maas, 1893 (sens. ampl.).

Gonads not localized as in Homceonema and Rhopalonema, but developed diffusely over

the radial-canals. Four lips.
HALICREAS, Fewkes, i$82 = Halicreas + Haliscera, Vanhoffen, 1902

= Isonema (in part), Maas, 1906.

The mouth is a simple round opening, without four lips. (In all known species the
radial-canals and ring-canals are very broad and flat.) Wart-like protuberances
may be present upon the sides of the exumbrella. Radial tentacles large, interradial,
small. Tentacles arise in a single row.
BOTRYNEMA, Browne, 1908.

Similar to Halicreas, but the tentacles are grouped in linear clusters in a single row

around the bell-margin.
CROSSOTA, Vanhoffen, 1902.

The tentacles arise in several rows from the bell-margin. Mouth with four lips.

Subfamily No. 2, AGLAURINjE.

Stomach mounted upon a peduncle.
AGLAURA, Peron and Lesueur, 1809.

Eight gonads upon the peduncle above the stomach. Sexes separate. Development direct.
AGLAmHA = /fglantha + 4glisi:ra, Haeckel, 1879.

Eight gonads upon the subumbrella, or at the turning points of the eight radial-canals

between the peduncle and the subumbrella. Sexes separate.
AMPHOGONA, Browne, 1904.

Similar to Aglantha, but medusa is bisexual, four of gonads being male, and four female.
STAURAGLAURA, Haeckel, 1879.

Four gonads, one upon each alternate radial-canal.
PERSA, McCrady, 1857.

Two gonads on two of the radial-canals, 180° apart. The six other radial-canals are sterile.

Family No. 6, GERYONIDjE.

Trachymedusae with four or six radial-canals upon which the flat, expanded, leaf-like
gonads are developed. Stomach mounted upon a gelatinous peduncle. The ring-canal
gives rise to blindly-ending centripetal canals.
LIRIOPE, Lesson, i%^ = Linantha -\-Lirwpc + Glossocodon -\-Glossoconus, Haeckel, 1879.

Four radial-canals. Four gonads. Four lips. With four primitive, solid, radial, and four
solid interradial, and four hollow, flexible, radially-placed tentacles; all twelve of
which may be found upon the medusa at one and the same time. Development
direct through a free-floating, actinula-like larva.
GERYONIA, Peron and Lesueur, i8og = Geryones + Geryonia + Carmaris

+ Carmarina, Haeckel, 1879.
Similar to Liriope, but with six radial-canals, six gonads, six lips, etc.; instead of four.

NARCOMEDUS.E. 15

Order No. 4. NARCOMEDUS^ Haeckel, 1879.

Veiled medusae with bell-margin cleft into intertentacular lappets. With free lithocyst-
clubs, containing concretions of entodermal origin. These medusx develop from actmula
larvae either directly or by budding. The bell grows outward from the sides ot the body of the
actinula, or the medusa-bud, leaving the tentacles stranded in the partially closed-over clefts
between the lappets of the bell. The Narcomedusae are thus medusiform, actinula-like
animals, the bell of which is not homologous with that of the Anthomedusae or Leptomedusae.

Family No. i, SOLMARIDjC.

Narcomedusae in which the outer margin of the stomach is plain, entire, and without
peripheral stomach-pouches. Saccules may, however, arise from the subumbrella floor of
the stomach.

o/)'fo//>a (young) + Sohnaris, Haeckel, 1879.
Without subumbrella saccules. Gonad is a simple annulus in ectoderm of subumbrella

floor of stomach.
PEGAJXTBA=Pegasia+Polyxenia+Pegantha+SoImoneta (in part), Haeckel, 1879.

With out-pocketings on the subumbrella floor of the stomach. The gonads are developed
in these subumbrella saccules.

Family No. 2, jEGINID^E, sens. ampl.

Narcomedusae in which the central stomach gives rise to simple or cleft marginal out-
pocketings in the radii of the tentacles.

CUNANTHA, Haeckel, 1879.

Four tentacles. Four peronial strands in the tentacular radii. Four simple, uncleft,
peripheral stomach-pouches in the radii of the tentacles. This "genus" is prob-
ably only a young stage of JEgina.
JEciNA, Eschscholtz, i$2g = Cunarcha + £gina + SolmunJus, Haeckel, 1879.

Four tentacles. Four peronial strands. Four cleft ( = eight peripheral) stomach-pouches,
outer margins of which may be still further divided.

SoLMUNDELl.A=dlgi>iella + SolmunJella, Haeckel, 1879.

Two tentacles. Four peronial strands. Four cleft ( = eight peripheral) stomach-pouches.
An apical (exumbrella) sense-organ is present in larva, but does not persist in adult.
Derived from jEgina by the disappearance of half ot its tentacles.

HYDROCTENA, DawydofF, 1903.

Two tentacles. No peronial strands. Two simple, uncleft stomach-pouches in the
tentacular radii. There is an apical (exumbrella) sense-organ consisting in a
ciliated pit containing two lithocyst-clubs. A median axial canal extends upward
from the stomach to the bottom of the sensory pit.

CUNOCTANTHA, Haeckel, 1879.

Eight tentacles. Eight peronial strands. Eight simple, uncleft stomach-pouches in the
tentacular radii.

unoctona + ^Eginura, Haeckel, 1879.

Eight tentacles. Eight peronial strands. Eight cleft ( = sixteen peripheral) stomach-
pouches. The outer margins of these pouches may be still further divided so as to
give thirty-two marginal pouches.

S, Brandt, 1835.

Four tentacles. Eight peronial strands. Eight cleft ( = sixteen peripheral) stomach-
pouches. Derived from jEginura by the disappearance of half of its tentacles.

CUNINA, Eschscholtz, 1829.

Nine or more tentacles, and an equal number of peronial strands. Peripheral stomach-
pouches simple, uncleft and equal in number to the tentacles, in the radii of which
they are developed. With otoporpae above the sense-clubs.

16 MEDUS.E OF THE WORLD.

SOLMISSUS, Haeckel, 1879.

Similar to Cunina, but without otoporpae.

CvNlSSA = Cnnissa + ^EginoJorus, Haeckel, 1879.

Nine or more tentacles, and the same number of peronial strands. Peripheral stomach-
lobes twice as numerous as the tentacles, being cleft in the tentacular radii.

jEciNODiscus, Haeckel, 1879.

Eight tentacles, sixteen peronial strands. Sixteen cleft ( = thirty-two peripheral) stomach-
pouches.

MEDUS/E MILLEPORIN^;.

Under this heading we may place the degenerate, free-swimming medusae of
Millepora. They have no velum and are thus separated from the veiled medusae
or Craspedotae. Not only is the velum absent, but the medusa is also devoid of a
peripheral canal system and of marginal tentacles.

The medusae Milleponna? and Craspedotae are doubtless derived from a
common ancestral phylum, but have departed widely, one from the other, so that
the Craspedotae are constantly characterized by a diaphragm-like membrane, or
velum, which partially closes the opening of the bell-cavity at the tentacular margin;
whereas this structure is absent in the Milleporinae.

The only known forms of Medusae Milleporinae are those of Millepora.

Millepora alcicornis (Medusa).

Millepora (medusa*), HICKSON, 1900, Proc. Roy. Soc. London, vol. 66, p. 3, figs. i-io. — DUERDEN, 1899, Journal of the Institute
of Jamaica, March, 1899. — HICKSON, 1906, Cambridge Natural History, vol. I, Coelenterata, p. 259, fig. 129.

Each medusa lies in an ampulla, or cavity, of the corallum, and is attached by a narrow
stalk to the innermost part of the wall of the cavity. The bell is 0.4 to 0.6 mm. in diameter
and is devoid of radial or circular canals or velum. It consists of a median lamella of entoderm
covered on the exumbrella and subumbrella sides by an ectodermal epithelium. There are
no tentacles, but instead there are 4 or 5 swollen masses of nematocysts 90° or 72° apart, near
the bell-margin, but projecting from the sides of the exumbrella. The manubrium is greatly
swollen, and fills the greater part of the bell-cavity. A mouth may (or may not ?) be present.
There are I to 5, usually 3 or 4, large ova in the ectoderm ( ?) of the manubrium. The central
entodermal cavity of the manubrium consists of an axial chamber which often gives rise to 4
perradial pouches which project into the entoderm of the manubrium, and end blindly. In
some medusae these pouches do not exist, in other specimens there are but 2 or 3, but probably
the most common condition is that of 4 pouches 90° apart. The fully-developed ova occupy
positions alternating with the pouches. Dr. Duerden observed that these medusae are set
free and swim slowly about with infrequent feeble pulsations. Soon after liberation the ova
begin to be discharged into the water and the medusa dies after 5 or 6 hours of life, having
discharged all of its eggs.

The Pacific millepores also produce medusae (see Hickson, S. J., 1891, Quart. Journ.
Micros. Sci., 1898; Proc. Zool. Soc. London; and Philosoph. Trans. Roy. Soc. London, B,
vol. 179).

OKOKIf A.Vmn.MKDI S.K. 17

VEILED MEDUS/E.
MEDUSAE CRASPEDOT^E Gegenbaur, 1856.

Cr\ftacar/iir, ESC'HSCHOLTZ, 1829, Syst. drr Acalc'phac.

Gymnopthalmeet FORBES, 184^, British Naked-eyed MeJuvr.

CnispeJala, GEC-.ENBAI'R, 1856, Zeit. fiir wisscn. Zool., Bd. X, p. 217.

Craspcdotf, HAECKEL, 1879, Syst. der Medusen. — MAAS, 1X9}, Ergi-h. ilcr Plankton-Expedition] Bd. 2, K.c.

CHARACTERS OK THK VKII.ED MEDUSJE.

Medusae with a velum or diaphragm which partially closes the marginal opening of the
hell-cavity; with ectodermal gonads (i. e., the sexual products u'lifn ripe are commonly found
in the ectoderm). Without gastric filaments, \\irh a douhle marginal nerve-ring, one ahmi-
and one below the velum. Development either direct or through alternation of generations
from hydroids.

Order ANTHOMEDUS^E Haeckel, 1879.

Oceanitit (in part), GEUENBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, pp. 218, 219.

Tubularix (in part), ACASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 337.

AnthomeJustr, HAECKEI., 1879, System der Medusen, p. 3. — VANHOFFEN, 1891, Zoologischer An/eiRtr, Bd. 14, pp. 439, 442.—
MAAS, 1905, Craspedoten Medusen der Siboga Expedition, p. 5; 1897, Mem. M. C. Z. at Harvard Coll., vol. 23, p. 9. —
HARTLAUB, 1892, Nachrichten kgl. Gesell. \Vissenschaftcn Gottingen, p. 17; 1897, Nordisches Plankton, Nr. 12, p. 4. —
HARGITT, 190^, Bull. U. S. Bureau of Fisheries, vol. 24, p. 29.

CHARACTERS OF THE ANTHOMEDUSyE.

Hvdromedusae in which the gonads are contained within the ectoderm of the manubnum.
There are no marginal otocysts or sensory clubs. The hydroids are of the Tubulanan order.
We may distinguish three families:

(1) CodoniJif with simple tentacles, and with the gonad in the form of one or more rings
encircling the manubrium. No oral tentacles.

(2) CladonemiJtr with feathered or branched marginal tentacles. Gonads ring-like, or
more or less separated. With or without oral tentacles.

(3) OceaniJir with gonads confined to the interradial or adradial sides of the manuhnum
and separated one from another in the meridians of the principal radii. With or without
oral tentacles. With unhranched marginal tentacles.

The bell in the Anthomedusae is without marginal lappets, and usually dome-shaped.
The tentacles arise from the bell-margin, not from the sides of the bell. The velum is simple
and annular, and is provided with circular muscles serving to produce the periodic con-
tractions of the bell. The hell-margin is simple and entire, and there are no otocysts, otohth-
clubs, or club-shaped sense-organs. Marginal cirri rarely present. In the young medusa the
exumbrella is more or less besprinkled with nematocysts, but these usually disappear or
become less conspicuous in the adult; although definite nematocyst tracts often persist over
the exumbrella, this being far more commonly seen in Anthomedusas than in other orders of
Hydromedusae.

The tentacles usually arise singly from the bell-margin, and are usually simple and
unbranched. Ectodermal ocelli are often found upon the tentacle-bulbs, and Romanes
showed that in Sarsia these subserved a visual function, and that the medusa was strongly
attracted by rays between the red and violet. The main shafts of the tentacles are thickly
covered with nematocysts, which may be mounted upon filaments or converted into adhesive
organs, as in the Cladonemidae. The tentacle-bulbs are hollow, and connected with the
gastrovascular system ot the medusa.

The radial-canals are usually simple, but in the Tiarinae they often give rise to short,
blindly-ending diverticula, which may be glandular in function, (ilandular swellings are
found in the walls of the radial-canals of L\tnn<ir,-ii iil,-\iinJri, and probably in those of /)y-
inorpliosa dubia and Sliikbi-ria hnltcrtita. A ring-canal is present in all genera excepting

18 MEDUS.E OF THE WORLD.

Pachycordyle and the Williadi, where it has become filled with a more or less solid core of
entoderm cells. The ring-canal is usually simple, and rarely gives rise to blindly-ending
diverticula.

The proboscis, or manubrium, is usually flask-shaped, and the mouth in most of the
Codonidae is a simple round, or cruciform, opening, but in the Oceamdae it is surrounded by
lips which are more or less folded or crenated. Oral tentacles are found in the Margelmae,
and in some Cladonemidae. The stomach is often mounted upon a solid, conical peduncle,
but in Turritopsis the peduncle is made up of large, highly-vacuolated, entodermal cells which
constitute the walls of the proximal parts of the 4 radial-canals. The medusae are carnivorous,
feeding upon Crustacea, fishes, and other medusae or Siphonophorae.

In the Codonidae the gonad encircles the manubrium in a ring-like manner, the mature
genital products being found in the ectoderm on the sides of the stomach. In the genus
Slabbena there are 2 or more of these genital rings. In the Oceamdae, however, the gonads
are restricted to the interradial, or adradial, sides of the stomach. Haeckel, 1879, believed
that they were radial in position, corresponding with the radial-canals, but VanhofFen, 1891,
showed that this was an error, for they are commonly interradial, or on both sides of each
radial line. In Eleutheria, according to Hartlaub, and possibly in other Cladonemidae, the
genital products are developed in the ectoderm of a peculiar brood-pouch, which is not
connected with the stomach, but is invaginated from the general ectodermal wall of the
subumbrella. In the Williadi and in Nemopsis the radial corners of the stomach extend
outward along the radial-canals, and the gonads develop upon the sides of these pouches.
Eleutheria is successively hermaphroditic, either sex preceding, but in all other Anthomedusae
the sexes are separate. The mature genital products are usually found in the ectoderm,
although they often originate in the entoderm. The eggs are cast out into the water by the
breaking down of the ectodermal walls ot the manubrium, but in some species of Bougatnvillia,
or in Margelopsis, or Hybocodon, the larvae may be retained until they have passed into the
planula or even into the actinula stage. None of the Anthomedusae are known to develop
directly from the egg into medusae, but the hydroid stages of many genera remain unknown.
Wherever the sexual development is known it is through hydroids of the Tubulanan order, in
which the medusae bud out separately and are not protected in special capsules or sporangia.

Asexual development of medusae is found in Codonidae, in medusae of the genera Hv/>o-
codon, Slabberia, and Sarsia, and in the Oceanidae in the genera Cytieis, Dysmorphosa, Bou-
gainvillia, Rtithkea, and in the Williadi. In some of the Williadi the medusa-buds are borne
upon stolons which arise either from the radial corners of the stomach, or from the forks of
the radial-canals. In the case of Sarsia and Hybocodon the budding medusae develop as in
hydroids, the entoderm of the parent medusa forming the entoderm of the budded medusa,
and the ectoderm of the bud being formed from the ectoderm of the parent. In Rathkea,
Chun, 1895, discovered that the budding medusa is formed out of the ectoderm of the parent,
although a connection is finally established between the entoderm of the bud and that of the
parent a short time before the hud is set free. In Bougainvillia tuobe, on the other hand,
I find that the budding medusa is formed entirely from the ectoderm of the parent, no con-
nection being established between the entoderm of the bud and that of the parent. In the
genus Niobia the tentacle-bulbs develop into medusae and are set free.

The majority of the Anthomedusae undergo a considerable development while swimming
freely in the water. In some cases, however, the medusae may, at times, become mature and
even discharge their genital products while still attached to the hydroid. This is seen in
Pennaria, Podocoryne, Sarsia, and Stylactis. Among the most short-lived medusae are those
of Pachycordyle, which have neither tentacles, marginal sense-organs, nor radial nor circular
canals.

All of the Anthomedusae are inhabitants of salt water. They are rare in the open ocean
far from land, but are abundant along coasts, especially along continental shores, and many
of them thrive in harbors where the water may be more or less brackish. They appear to be
the simplest, and phylogenetically the oldest, of the Hydromedusae.

ORDER ANTHOMEDlS.i:. 19

Family CODONIDjC Haeckel, 1879, sens, amend.

Sarsiadat (in part), Founts, 1848, British Naked-eyed Medusa;, p. 55.

Sarsiada+ Tubularidtf, McCRADY, 1857, Gymn. Charleston Harbor, p. 21.

Sarsiadtf (in part)-t- Tubularidx+ Pennaridj: (in part), AGASSIZ, L., 1862, Cont. Nat. Hist. (-'. S., vol. 4, pp. ^9, 542.

SarsiadtE+ Orthocorynidx + Tubulands + Pennanda, AGASSIZ, A., 1865, North Amer. Acalephx, pp. 175, 183, |S6, 189.

Codonidz, HAECKEL, 1879, Syst. der Medusen, p. 9. — VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14^.442. — HARGITT, 1904, Bull.

Bureau of Fisheries U.S., vol. 24, p. 29. — HARTLAL-B, 1907, Nordisches Plankton, Nr. 12, pp. 5, 6.
Codonina+ Cor\nid<z + Tubularidtt, VON LENDENFELD, 1884, Zool. Anzeiger, Bd. 7, pp. 446, 447.

FAMILY CHARACTERS.

Anthomedusae in which the gonad is ring-like and encircles the manubrium. Mouth with-
out oral tentacles or prominent lips. The 4 to 6 radial-canals are simple and unhranched.
\Yhen ocelli are present they are found upon the outer sides of the tentacle bulbs. There
are no marginal otocysts. The tentacles are neither branched nor feathered.

Medusae of the genera Sarsia, Steenstrupia, Ectopleura, Pennaria, Amalthtfa, Trichorhiza,
Hyhocodon, H yJnc/ithys, Coryrutts, and Margt-lopsis are known to develop asexually through
alternation of generations from Tubulanan hydroids. Direct development of medusae from
the egg is unknown in this family, although medusae are budded asexually from the manubrium
of some species of Sarsia, from the basal bulbs of the tentacles of HybocoJon, and from those
of Sarsia. Actinula larvae are set free from the manubrium of Margelopsis and Hybocodon.

Haeckel, 1879, considered the presence or absence of an apical projection upon the bell,
and the presence or absence of an axial canal extending upward from the stomach into this
apex, to be a criterion for the separation of genera. It should be borne in mind, however,
as was pointed out by VanhofFen, 1891 (Zool. Anzeiger, pp. 439-446), that the young medusae
often lack an apical projection and axial canal, whereas the mature individuals possess these
characters. They are also highly variable in development. We have therefore considered
these characters to be of specific but not generic value.

A natural classification of the Codonidae can not be based upon the characters of the
medusae alone, for the medusae of Pcnnana and Amaltheea are similar, while their hydroids
are distinct, the former being Pcnnana and the latter Carymorpha, and an equally remarkable
condition is presented by hydroids belonging to the two genera Syncoryne and StaunJi/i,
both ot which give rise to medusae belonging to the genus Sarsia. This peculiar case is still
further complicated by the fact that at least one species of StauriJia hydroid gives rise to a
ClaJonema medusa.

In HyJrichthys we meet with an extraordinary case of parasitism or commensalism of
the hydroid, and in Margelopsis we find a free-floating hydranth, recalling the hypothetical
ancestral form of the Siphonophorae. Margclnpsis is also interesting in that its medusae form,
apparently, a connecting link between the Codonidae and the Bougamvilleae, although they
are more closely related to the Codonidae than to the last-named family. Indeed their
resemblances to the Bougamvilleae are probably due to mere parallelism and not to blood-
relationship.

The Codonidae may conveniently be divided into two subfamilies, as follows:

(1) Srirsi/inn, with simple, unbranchtd, marginal tentacles, which arise singly from the bell-margin.

One or all of the tentacles may he degenerate or absent.

(2) Margelopstna, with marginal tentacles grouped into clusters.

20 MEDUS.E OF THE WORLD.

The following table defines the characters of the genera ot Codomdae.

SUBFAMILY No. i. SARSIAN^E.

With simple marginal tentacles which arise singly from bell-margin. One or all of the tentacles may be rudimentary or absent.

Tribe I. ANEMIDI.

Without tentacles or chymifcrous canals.

Pachycordyle, WEISMANN, 1883; HARCJITT, \^o^=Parvanenius ( ?) MAYER, 1904.

Medusa without tentacles, marginal sense-organs, radial-canals or circular canal. Hydroid: Pachycordyle ', WEISMANN.

Tribe II. AMALTHIADI.

The hydroid is Corymorpha. Medusae have 4 radially placed, rudimentary tentacles and 4 radial-canals.
Amalthtza, SCHMIDT, 1854. M.edus*= Amafthaa, HAECKEL.

With 4 rudimentary tentacles, 4 radial-canals, and ring-canal. Genital products on manubrium. Hydroid: Corvmorpha.

Tribe III. PENNARIDI.

Medusae similar to those of Tribe II, but hydroid is Pcnnaria.
Pennaria, OKEN, 1815; GOLDFUSS, 1820.

With 4 rudimentary tentacles, 4 radial-canals, and ring-canal. Genital products within manubrium. Hydroid: Pennaria.

Tribe IV. PSEUDOPENNARIDI.

Medusa similar to those of Tribes II and III, but hydroid is Trichorhiza.
Trick or kiza, RUSSELL, 1906.

Medusa similar to Pcnnaria, but hydroid is Trichorhiza.

Tribe V. EUPHYSIDI.

With small or rudimentary tentacles situated at bases of all but one radial-canal, and with one or more well-developed tentacles
situated at base of remaining radial-canal. Bell may or may not possess an apical projection, or an axial canal extending
into it from stomach-cavity.
Sleenstrupia, FORBES, 1846= Euph \sa, FORBES, 1848; Eufthysora, MAAS, 1905.

The bell is radially symmetrical and the 4 radial-canals are all of equal length, i long and 3 short tentacles. Hydroid:

Corymorpha.
ffybocodon, L. AGASSIZ, 1862= Amphicodon, HAECKEL, 1879.

The bell is asymmetrical. One of the 4 radial-canals is long, the one opposed to it short, and the two other canals of inter-
mediate length. One long tentacle arises from the base of the long radial-canal, while the other 3 tentacles are short
In old medusae of Hybocodon one is apt to find 2 or more tentacles at the base of the long radial-canal, and the medusa,
is in the Amphicodon stage. Hydroid: Hybocodon.
Microcampanay FEWKES, 1889.

With 6 radial-canals and 6 tentacles, one of which is well-developed and 5 arc rudimentary. Hydroid unknown.

Tribe VI. DICODONIDI.

With 2 well-developed, diametrically opposite tentacles, and 2 rudimentary tentacles. With or without an apical projection

and with or without axial canal extending upward from the stomach into apex of bell.
Dicoeloniums HAECKEL, 1879.

With 4 simple radial-canals and 4 radially placed tentacles; 2 of the tentacles are long, and 2 short. Gonad is ring like
and surrounds stomach. No lines of nettling cells over exumbrella.

Tribe VII. SARSIADI.

Bell radially symmetrical. With 4 equally-developed, unbranched, marginal tentacles. With or without an apical projection,
and with or without an axial canal extending upward from stomach into apex of bell, (i) The manubrium is encircled
by a single ring-like gonad. (2) With 2 or more ring-like gonads upon the manubrium.
Sarsia, LESSON, i$4$ = S\ndict\ont A. AGASSIZ, 1862; Codoniumt HAECKEL, 1879.

Manubrium slender and tubular. Each tentacle-bulb bears an abaxial, ectodermal ocellus. Hydroid: S \ncoryne.

Also Staiindta.
Hydrichthys, FEWKES, 1888.

Medusa similar to Sarsia, but without ocelli upon the tentacle-bulbs. Hydroid: Hvdrichthys.
Eneodonium, HARTLAUB, 1907.

Medusa similar to Sarsia, but stomach is mounted upon a gelatinous peduncle, and tentacles end each in a knob.
Ectopleura, L. AGASSIZ, 1862.

8 longitudinal lines of nematocysts extend from bases of tentacles to apex of bell. Hydroid: Ecto pleura.
Corynitis, McCRAov, 1857.

Manubrium cruciform in cross-section, and bound to the 4 radial-canals by hollow mesenteries. 4 radially placed marginal
tentacles which are covered with wart-like clusters of nematocysts. Their basal bulbs bear ocelli. Hydroid: Syncoryne.
Slabber ia, FORBES, 1846= Difwrena, McCRADY, 1857. Dif>urena + Bathvcodon, HAECKEL.

Similar to Sarsia, but with two or more ring-like gonads upon the manubrium. Hydroid: S\ncor\ne.

SUBFAMILY No. 2. MARGELOPSINJE.

With 4 radiallv-placed clusters of marginal tentacles. No oral tentacles. Gonads ring-like, and encircling the manubrium.
MargelopsiSy HARTLAUB, 1897.

Marginal tentacles are grouped in 4 radially-situated clusters. Gonad ring-like, and surrounds manubrium. No oral
tentacles, mouth is a simple round opening. Medusae develop by budding from a free-floating hydranth, A/drgr/o/fj/i.
This interesting genus recalls the Bougainvillidi in its radially situated clusters of marginal tentacles. It resembles the
Codonidts in its ring-like gonad, and in the absence of prominent lips and oral tentacles.
Pelagohydra, DENDY, 1903.

Medusa similar to Margelopsi* but hydroid is Pelagohydra, and differs from hydroid of Margelopsn in that tentacles arise
at irregular intervals from the sides of the hydranth and are not arranged in definite verticils as in Margelopsis.
Hydroid is pelagic.

PLATE i.

Fig. I. PaehycorJyle degeneratus, male. Nassau Harbor, Bahama Islands,
July 19, 1903.

Fig. 2. Budding hydranth of Pennaria tiarella with female medusa-bud
still attached. Agassiz Laboratory, Newport, Rhode Island,
July 5, 1895. Found attached to eel grass in shallow water.

Fig. 3. A recently liberated male medusa of Pennaria tiarella. Agassiz
Laboratory, Newport, Rhode Island, July, 1895.

Fig. 4. Hydroid stock of Pennaria tiarella, natural size. Agassiz Labora-
tory, Newport, Rhode Island, July, 1895.

Fig. 5. Pennaria tiarella. From the coral reef at Tortugas, Florida, May 7,
1905.

fig. 6. Steenstrupia vtrgulata. From a drawing made from life by Dr.
Alexander Agassiz at Nahant, Massachusetts, August 21, 1862.

Fig. 7. Steenstrupia rubra, male. Oregon Inlet, Pamlico Sound, North
Carolina, November 12, 1904.

Fig. 8. HyhoeoJon forbesii. Tortugas, Florida, May 25, 1905.

Fig. 6, from a drawing by Dr. Alexander Agassiz
Figs, i to 5, 7, 8, from life, by the author.

AXTHOMKIHS K I'A( IIV( -iililiVLK. '_' I

DESCRIPTION OF GENERA AND SPECIES OF ANTHOMEDUSjE.

Genus PACHYCORDYLE Weismann, 1883.

Pachycordylt, WEISMANN, 1883, Entstehung Seiualzellen Hydromedusen, pp. 87, 217. HAKI.II r, 1404, Mitth. Zool. Station

Neapel, BJ. |6, p. 553.
Paminrmus, MAYER, 1904, Mem. Nat. Sci. Museum Brooklyn Institute Arts and Sti., v.il. i, No. i, p. 6.

GENERIC CHARACTERS.

Codonidae without tentacles, radial-canals, or circular vessel. Manubrium surrounded
by a ring-like gonad. The hydroid-stock is Pachycordyle.

Except the medusas of Millepora, these are the most degenerate and short-lived of the
free-swimming Hydromedusae. They are even more degenerate than are the medusae of
Corymorpha and Pennaria, and may be compared with Eucopflla (R. von Lendenfeld, 1883,
Zeit. fur wissen. Zool., Bd. 38, p. 497) and Agastra (C. Hartlaub, 1897, Wissen. Meeresunter-
such. Biologisch. Anstalt auf Helgoland, Neue Folge, Bd. 2, Heft i, Abt. 2, p. 504, taf. 22,
figs. 5, 8 10). In these last-named medusae we find neither manubrium nor marginal tentacles,
hut there are 8 otocysts and a velum, and the radial and circular vessels are well developed.

Pachycordyle weismanni Hargitt.

Pachvcorii\lf lutnmannt, HAROITT, 1904, Mittli. Zool. Station Neapel, Bd. 16, Heft 4, p. 553, plate 21, figs. 1-8.

( ?) Pachyconlyle napolilana, WEISMANN, 1883, Die Entstehung der Seiualzellen bei den Hvdromedusen, Jena, pp. 87, 217.

Medusa pyriform, 2 mm. high, 1.3 mm. wide. Tentacles and marginal sense-organs
lacking. No radial-canals. Ring-canal a mere fissure with vestiges of an entodermal linin»
near the margin. Velum narrow, with a small opening. Manubrium large, conical, and with-
out a peduncle. Ripe ova are in the entoderm, and are discharged very soon after medusa is
liberated. Mouth lacking. Manubrium orange or dark-brown, other parts colorless. The
medusa swims with a short, jerky motion, but lives only one or two hours.

The hydroid is found in the Bay of Naples growing upon the shell of Fusus rostratits.
The colony arises from a delicate, reticulated hydrorhiza. Hydrocaulus sparingly branched,
3 to 8 mm. high. Perisarc dull yellowish-brown, not extending beyond base of the hydranth.
Hydranths club-shaped, with subconical hypostome. 8 to 16 irregularly arranged filiform ten-
tacles, delicate and thread-like when expanded. Body of hydranth orange or reddish,
hypostome whitish. Not more than 2 or 3 medusa-buds are borne on the side of the stem
of each budding polyp. Occasionally the medusa-buds develop on the side branches, more
commonly on the main stems. The ova originate in the entoderm where they remain until
discharged into the water from the manubrium of the medusa. This species may possibly be
the female torm of VVeismann's Pachycordyle napolitana. On Weismann's hydroid, how-
ever, there were no free medusae, only sessile gonophores. All of Weismann's specimens were
males, while Hargitt's were females, and it is possible, as Hargitt states, that the females
only give rise to free-swimming medusae.

Pachycordyle degeneratus.

Plate I, fig. I.

Paroanemu* degtneretuit MAVFR, 1904, Memoir-. N.it. Sn. Museum Rrooklvn Institute of Art^. and Sci., vol. i. No. i, p. fi, plate
',. "H- "•

Bell thin-walled, about 0.75 mm. high and 0.3 mm. in diameter. Bi-ll-walls quite rigid,
velum powerful and well developed. There are neither tentacles, radial-canals, circular
vessel, nor marginal sense-organs. Manubrium spindle-shaped, and about a third as lon»
as height of bell. Fluids within the stomach-cavity are maintained in rapid motion, appar-
ently through the action of cilia. Near aboral end of bell is a deep conical cicatrice which
apparently marks the place of last connection between the medusa and its hydroid stock.
Bell is translucent and milky in color, while manubrium is cream-colored. Only 5 specimens,
all males, were found in Nassau Harbor, Bahamas, on the nights of July 18 and 19, 1903.

22 MEDUS/E OF THE WORLD.

They swam actively in arcs of circles, but all died early in the morning although maintained
in large glass dishes filled with pure sea-water. They appeared to be mature, for sperm was
discharged constantly from the sides of the manubrium.

The hydroid is unknown, and possibly the medusa may not belong to the genus Pachy-
corJyle, though in the medusa stage it conforms with the generic character of the medusae
which are set free from Pachycordyle iveismanni of Naples.

Genus AMALTH^EA Schmidt, 1854.

Corvinorplia (hydroid), SARS, M., 1835, Beskriv og Jagttagelser, p. 7.

Corymorpha (medusa and hydroid), STEENSTRUP, 1854, Vidensk. Meddel. Nat. For. Kjobenhavn, p. 46.

Amahhxa (medusa), SCHMIDT, 1854, Handatlas vergl. Anatomie, p. 13. — HAECKEL, 1879, Syst. der Medusen, p. 38.

Amallhiza, ALLMAN, 1872, Monog. Tubularian Hydroids, p. 393. — LOMAN, 1889, Tijdschrift, Nederland. Dierk. Vereen, Ser. 1,

Deel. 2, p. 270.
Corymorpha, HARTLAUB, 1905, Zoolog. Jahrb., Abth. Syst., Suppl. 6, p. 543; 1907, Nordisches Plankton, Nr. 12, p. 75.

GENERIC CHARACTERS OF THE FREE MEDUSA.

Codonidae with 4 rudimentary tentacle-bulbs, 4 radial-canals, and a ring-canal. Hydroid
is Corymorpha, but distinguished by the character of its medusae. The majority of species
of Corymorpha hydroids do not give rise to free-swimming medusae. Hydroid of Arnalthtta
might therefore be called Amalthaa. The medusae of Amalthcra resemble those of Pennaria,
but the manubrium is usually longer, and extends beyond the velar opening.

Hartlaub, 1907, includes the medusae of Steenstrufia, Euphysa, and Amalthaa in the
genus Corymorpha, for they all arise from identical hydroids; while the medusae differ as
follows: Steenstrufia, 3 short, and I long tentacle, and with an apical projection and axial
canal above the stomach. Euphysa, similar to Steenstrufia, but without an apical projec-
tion or axial canal. Amalthxa with 4 rudimentary tentacles.

Amalthrea sarsii Allman (Medusa only).

Corymorpha sarsii, STEENSTRUP, 1854, Vidensk. Meddel. Naturhist. For. Kjobenhavn, p. 48. — BONNEVIK, 1898, Zeit.fiir wissen.

Zool., Bd. 43, p. 476. — HARTLAUR, 1907, Nordisches Plankton, Nr. 12, p. 86, fig. 82.

Amatthtea sarsi:, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 393. — HAECKEL, 1879, Syst. der Medusen, p. 38.
( ?) Arnaltluea amtebigera, HAFCKEL, Ibid., p. 38, taf. I, figs. 10, ll.

It is not certain that this hydroid gives rise to free-swimming medusae, though when arti-
ficially set free the medusae swim about actively.

The bell of the attached medusa-bud is elongate with a subconical basal apex and with
fairly thick bell-walls. It is 4 mm. high. There are usually 4 equally developed rudimentary
tentacles, but occasionally one of the tentacles is longer than the others. Velum well developed.
4 straight, slender radial-canals. Manubrium spindle-shaped. Mouth a round opening
studded with nematocysts. In the male the manubrium projects one-third its length beyond
velar opening, but in the female it is shorter and the mouth is at the level of velar opening.
Ova are few in number. When immature they are amoeboid, but when older they become
spherical and project over the surface of the gonad, attached by short pedicels. The gonad
encircles the entire manubrium. Manubrium straw-yellow, tentacle-bulbs light-red. Found
oft coast of Norway. Hydroid: Corymorpha sarsn.

Amalthaea uvifera Schmidt.

Amalthira uvifera, SCHMIDT, 1854, Nordisches Plankton, Nr. 12, p. 88, fig. 83.

Corymorpha uvifera, HARTLAUB, 1907, Handatlas der vergleich. Anatomie, p. 13, taf. 9, fig. 2.

This medusiform gonophore is not known to be set free. It is borne upon branched
stolons which arise from the sides of the polypite above the basal circlet of long tentacles.
Each stolon is tree-like and gives rise to 30 to 40 grape-like medusa-buds, so that each
hydroid gives rise to more than 100 buds in various stages of development.

In the medusa-bud (when about to be set free ?) there are 4 large globular, rudimentary
tentacle-bulbs. The bell is higher than a hemisphere, with walls thin at apex. Manubrium

ANTHOMKDUS.K— AMAI.TII.KA, I'KN \.\HI.\. -'.',

spindle-shaped, as long as the depth of the bell-cavity. The hydroid is C.or\in<n [>hn uvifern,
found at Loppen Island, ahout 10 miles from Hammerfest, at a depth of about I fathom.

Amalthaea vardoensis Loman.
Amahh<ta vardiitnsis, LOMAN, 1889, Tijdschr., Nederland. Dierk. Ver., Ser. 2, Dcel. 2, p. 271, teit-fig. 5, taf. 13, figs. 1-9, 15.

Hydroid about 50 mm. high. The thin, transparent perisarc extends only over lower halt
of stem. The terminal polypite is sharply set off from the stem by a constriction at its base.
Nearly 50 large tentacles in basal circlet at wide base of polypite. Over 100 very short tapering
oral tentacles in 7 to 9 circlets. 16 to 20 peduncles arise in a circlet from the sides of the
polypite between the basal and oral tentacles. Each peduncle bears a number of medusa-
buds, which have 4 radial-canals and 4 equally developed, very short tentacle-bulbs. The
hydroid is translucent rose-colored. Found in Busse Sound at Vardri, 71° N. lat., Norway.
For details of histology, see Loman.

Amalthaea (?) Hybocodon(?) januarii Steenstrup.

Corymorpha januarii, STEENSTRUP, 1854, Vidensk. Meddcl. Nat. For. Kjbbenhavn, p. 46.— SARS, 1861, Annals and Mag. Nat.

Hist., vol. 8, p. 356.
Amalthtfa januarii, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 394. — HAECKEL, 1879, Syst. der Mi-JuM-n, p. 39.

Described by Steenstrup from a single imperfect hydroid found in the harbor of Rio
Janeiro, Brazil. The hydrocaulus is about 150 mm. long and 8 mm. wide. About 80 tentacles
in the proximal circlet, and these are about 50 mm. long. The oral circlet was imperfect
and can not be described. There were about 40 branched peduncles above the basal circlet of
tentacles. These bear numerous medusa-buds having 4 equal tentacle-bulbs, but oblique
margins. The free medusae are unknown.

Genus PENNARIA Oken, 1815.

Pennaria (in part), OKEN, 1815, Lehrbuch der Naturgesch., Bd. I, p. 94.

Pennaria (hydroid), GOLDFUSS, 1820, Handbuch der Zoologic, p. 89.

Pennarin (medusa), MI/CRADY, 1857, Gymn. Charleston Harbor, p. 50.

Pennaria + Halocordyle, ALLMAN, 1871, Monog. Tubul. Hydr., pp. 363, 368.

Pennaria, AOASSIZ, L., 1861, Cont. Nat. Hist. U. S., vol. 4, p. 278. — AC;ASSIZ, A., 1865, North Amer. Acal., p. 187.— HARTLAUB,

1907, Nordisches Plankton, Nr. 12, p. 72. — PEEBLES, 1903, Archiv. Entwick.-mech., Bd. [4, p. 55 (regeneration).
Pennaria, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161 .--CLARKE, 1907, Mem.

Museum Comp. Zool. Harvard Coll., vol. 35, p. 6.
GlobicfpSf AC.ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344. — VANHOF* EN, 1891, Zool. An/eiger, p. 443. — HAECKF.I , 1879,

Syst. der Medusen, p. 39.

Globicfps (hydroid), AVERS, 1852, Proc. Boston Soc. Nat. Hist., vol. 4, p. 193.
Rucorvne (hydroid), LEIDV, 1855, Journ. Acad. Nat. Sci. Phila., ser. 2, vol. 3, part. 2, p. I VI- plate 10, figs. 1-5.

The type-species ot this genus is Pennaria ilistielia ot Goldfuss, 1820, from the Mediter-
ranean. Oken's "Pennaria" is wholly indefinite, including as it does Plumularia, Aglao-
phenia, etc.

(JKNERIC CHARACTERS.

Codonidae with 4 permanently rudimentary tentacles, which are reduced to mere basal
bulbs. The hydroid stock is a Pennaria.

Medusae of the genus Pennaria may become mature, and discharge their genital products
while still attached to the hydroid, and at the same time other individual medusae from the
same stock may be set free in an immature state. Generally, however, the genital products are
discharged a few hours after the medusae are set free into the water.

There is no generic difference between the medusae of Pennaria and medusae belonging
to the genus Amalthtsa, but their hydroids are different. The hydroid stock of Amalthaa
is Corvnifjrpha, and is closely related to the hvdroids of the medusa genera Hybocodon, St., n
strupia, and Ectopleura. The hydroid of the medusa genus Pennariti McCrady is Pennaria
Goldfuss (Globiceps Ayers). On account of this decided difference in their hydroids we
have separated Pennaria from Amaltheea = (Corymorpha\ for their apparent similarity is
only a case of parallelism.

24

MEDUSAE OF THE WORLD.

Pennaria disticha Goldfuss (European).

( ?) Pennaria marina, IMPERATO, 1599, Dell' historia naturale libri ventoto, Napoli, p. 747.

Pennaria cavoiini, SCHNEIDER, 1892, Jena. Zeitsch. Naturw., Bd. 20, p. 435, taf. 14, figs. 49-54 (histology).— Du PLESSIS, 188
Mittheil. Zool. Sta. Neapel, Bd. 2, p. 147. — PHTET, 1893, Revue Suisse Zool., tome i, p. 12, plate I, figs. 7-9.— WEI

, 1881,

is-

1903,
colonies).

colonies).
Pennaria disticha) BEDOT, 1901, Revue Suisse Zool., tome 9, p. 459; tome 13, 1905, p. 96 (citation of papers to 1850). — GOLD-

fucc iSii-i T-Ian^hiirh t\e*r 7.rwil . 11 Hn

FUSS, 1820, Handbuch der Zool., p. 89.
Pennaria symmetric a, CLARKE, 1879, Bull, Mus. Comp. Zool. at Harvard College, vol. 5, p. 240, plate I, figs. 2, 3.

Pennaria , PEEBLES, 1903, Archiv. Entwickelungs-mech., Bd. 14, p. 55, fig. (regeneration).

Sertularia fiennaria, DELLE, CHIAJE, 1841-44, Animali senza vertebra del Regno di Napnh, vol. 5, pp. 5, 17, plate 157, figs. 3,

14, 15; Ibid., 1822, tav. 43.

(tySertuIaria pennaria, LINN^US, 1758, Systema Naturae, edit. 10, p. 813.
Seriolara ftennaria, CAVOLINI, 1785, Mem. Polipi marini, Napoli, p. 134, plate 5.
Globiceps globator, HAECKEL, 1879, Syst. der Medusen, p. 40.
( ?) Medusa, Euphysa globaior, LEITKART, 1856, Archiv. fur Naturgesch., Bd. 22, p. 28, taf. 2, fig. 4.

Fin. i . — Pennaria disticha, from nature, by the author, from specimens collected by Dr. Lobianco in the Bay of Naples.
Ay hydrocaulus, natural size. B, terminal branches showing law of growth. C, terminal polypite. D and £, hydranths
showing completely-annul.ited pedicels.

ANTHOMKDUS.E — PENNARIA. I'.")

This hydroid is one of the earliest forms known, yet reliable information from the system-
atic standpoint is still incomplete. One of the best modern descriptions is that <>t Allman,
1872. I believe the American " Penmn-in tinn-lln" to be closely related to P. Jiiliilm.
The only difference appears to be that the terminal ramuli in P. disticha are ringed through-
out, whereas in P. tiarella they are usually ringed only at base and summit, although I have
seen an occasional one ringed throughout in the American hydroid. (See VV. S. Wallace,
1908, Year Book of the Carnegie Institution.)

In the European hydroid, P. Jisticha, the stems become about 150 to 175 mm. high. The
main stem is slightly zigzag and with a uniform growth-curve from base to summit. There are
about forty side branches, regularly alternate. Perisarc annulated at the origins of the branches
and on the ultimate hydranth-bearing ramuli. Hydranths flask-shaped, with a single verticil
of about 10 to 13 basal filiform tentacles each about I to 2 times as long as the body of the
hydranth, and each ending in a blunt, slightly swollen tip. About 20 short, stiff, knobbed
tentacles, irregularly arranged in 3 verticils, arise from the sides of the hydranth above the
basal verticil of tentacles. The medusa-buds are similar to those of Pennaria tiarella and
are without ectodermal ocelli upon their tentacle-bulbs. The medusa? usually wither upon
the hydroid without being set free, but this often occurs also in P. tiarella, especially in the
warm water of Florida.

Weismann, 1883, found that the germ-cells originate in the ectoderm of the inner layer
of the bell-nucleus and do not wander from their place of origin, becoming mature in the
free medusa, or when the medusa-bud is ripe.

Very elaborate studies upon regeneration, regulation, and restitution in injured colonies
were carried out by Cast and Godewski, 1903.

This hydroid is found in the Mediterranean. Pictet, 1893, found it at Amboina, Malay
Archipelago. It occurs at Naples, Italy, from May to November.

Pennaria tiarella McCrady (American).

Plate I, figs. 2-5.
REFERENCES TO THE AMERICAN HYDROID.

G/obiceps tiarella, AOASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344. — AVER, 1852, Proc. Boston Soc. Nat. Hist., vol. 4,
p. 193. — HAECKEL, 1879, Syst. der Mejusen, p. 39.

Pennarin tiarella, AGASSIZ, A., 1865, North. Amer. Acal.,p. 187, figs. 311-315. — NUTTING, 1901, Bull. U. S. Fish Commission
for 1899, vol. 19, pp. 337, 374, figs. 14, 83. — HARGITT, 1900, Amer. Naturalist, vol. 34, p. 387, plates 1-4, 36 figs.;
Bull, of the Bureau of Fisheries U. S., for 1904, vol. 24, p. 32, plate 3; 1904, Archiv. fur entwickrlungs-mech. organ-
ism, Bd. 18, p. 453, taf. 24-28; 1902, Amer. Naturalist, vol. 35, p. 31 1, figs. 8, 9; p. 597, fig. 36; 1899, Biol. Bulletin
Woods Hole, vol. i, p. 35-40, 6 figs, (grafting eiperiments); 1900, Science, New series, vol. 12, p. 340; and 1901,
Biol. Bulletin Woods Hole, vol. 2, p. 223 (variations of hydroids and medus.T.) — MCCRADY, 1857, Gymn. Charleston
Harbor, p. 51. — SMALLWOOD, 1899, American Naturalist, vol. 33, p. 861, 7 figs, (histology). — HARTLACB, 1907, Nor-
disches Plankton, Nr. 12, p. 72, figs. 68-70. — GOLDFARB, 1906, Journ. Experimental Zool., vol. 3, p. 148 (regeneration).

Pennaria liarclla=(P. svmmetrica, CLARKE), CONGDON, 1907, Proc. American Acad. Arts and Sci., vol. 42, p. 464.

Pennaria, j/>., THACHER, 1903. Biol. Bulletin, vol. 4, p. 96.

I'tnritiria gibbosa, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 278, 344; vol. 3, plate 15, figs. I, 2.

Halcord\Ie tiarella, ALLMAN, i87i,Monog. Tubul. Hydroiils, p. 369.

Kuforyne elegant, LEIDV, 1855, Marine Invert. N. J. and R. I., p. 4, plate 10, figs. 1-5.

THE AMERICAN PENNARIA.

Medusa. — The bell is about 2 mm. in height, and is ellipsoidal, being higher than it is
broad. Bell-walls very thin, flexible, and much distorted by the remarkably large ova, which
are situated within the ectoderm of the manubrium and often fill the entire cavity of the bell.
There are 4 small, rudimentary tentacle-bulbs, without ocelli, I at the base of each radial-
canal. Radial-canals straight and quite broad. Velum wide and powerful. Manubrium
of male more slender than that of female, the latter being usually greatly distended with 4 or
5 large ova. Several ova are often set free into the water before the medusa is liberated from
the hydroid stock. The medusa is commonly mature at the time of its liberation, and it is
probable that it does not usually survive for more than a few hours, though Dr. A. Agassiz
has maintained them alive for several weeks. The entoderm of the manubrium is rose-pink,
and there are a number of deep-pink blotches in the entoderm of each radial-canal. The
entoderm of the tentacle-bulbs is pearly-white, as are also the eggs within the manubrium.
These colors vary considerably in hue and may be almost white.

2fi

MEDTS.E (>K THE WOULD.

Hydroid. — The hydroid is very abundant upon eel-grass, seaweeds, stones, or wharves,
below low tide. The stems arise from a ramifying stolon. They attain a height of 100 to 125
mm. The main stems branch alternately, the longest and oldest side branches being found
nearest the base of the stock. The side branches give rise to simple or slightly branched

ramuli from their upper sides. These ultimate
ramuli are annulated at base with 5 to 6 rings, and
the main stem and primary branches are annulated
beyond each branch. The stems are covered with
a chitinous, horny perisarc. The polyp-heads are
flask-shaped, and the mouth is situated at the ex-
tremity of a long, slender, conical throat-tube.
There are 2 or 3 verticils of oral tentacles, each
row being composed of 5 to 7 short, stiff tentacles,
each terminating in a knob-shaped cluster of nem-
atocysts. Besides these there is a single circlet of
12 to 16 long, flexible tentacles near the proximal
base of the polypite. The medusa-buds develop
upon the sides of the polypites between the oral
and basal rows of tentacles. There are usually not
more than 2 or 3 buds upon each polypite at the
same time. The entoderm of this hydroid is white
to rose-colored, the ectoderm silvery and translu-
cent, and the perisarc horny-yellow to black.

This hydroid ranges from the West Indies and
Bermudas to the coast of Maine. Pentiaria gibbosa
L. Agassiz, of the Florida coast and West Indies,
is probably identical with P. tiarella, but the
medusae of the southern form wither on the stem
as do those of P. Jisticha. Indeed Agassiz's figure
in 1862, shows the ultimate pedicels of his P. gibbosa, from Key West, Florida, ringed
throughout as in the Mediterranean Pennaria disticha.

Hargitt, 1900, has made a study of the life-history and development of P. tiarella. Early
in the summer the hydroids are found growing on rock-weed, piles of docks, etc.; while late
in the summer they take to eel-grass. The early summer brood is not so bright in color, and
the medusae mature more slowly than in the late brood. The individual hydranths of the
late brood are, however, smaller than those of the early summer brood. The medusae dis-
charge their ova soon after liberation, and then die rapidly. The ova are 0.4 to 0.5 mm. in
diameter and are heavily laden with yolk. They are creamy-white to orange in color. The
cleavage is found by Hargitt to be subject to much individual variability, so that an extra-
ordinarily irregular, loosely connected mass of cells is formed, resembling the condition
described by Rittenhouse, 1907, in Turritopsis nutricula. No matter how irregularly shaped
the embryo may be, it results finally in the formation of a spherical solid morula, and this
soon changes into a pear-shaped, ciliated, planula larva. In 5 to 7 days after the beginning
of development, the larvae settle down and then develop into small monogastric hydroids
covered with ringed perisarc, and provided with whorls of tentacles as in the adult. Hargitt
finds that the removal of small parts of the segmenting eggs does not alter the future history
of development. The eggs may also be divided at the first or second cleavage and will still
develop into normal larvae. Moreover, Hargitt finds that single eggs sometimes separate
normally into two masses, each of which gives rise to a normal embryo. The irregularities
in cleavage observed by Hargitt have been seen also by Miss Cora J. Beckwith. She finds
that the segmentation is mytotic, not amitotic; the nuclear division constantly outnumbering
the cytoplasmic so that a syncytium is formed.

Hargitt, 1901, states that hydroid stocks of Pennaria which grow upon eel-grass in shallow
water are more pinnatified in their branching, and more highly colored than are those which
grow upon stones, etc., in deep water. He also finds that the medusae of the deep-water
hydroids are less active than are those developed upon the hydroids which grow in shallow

Flo. 2. — Southern variety of Pennaria tiarella. From
life, by the author, Tortugas, Florida, May
27, 1908.

Branch of hydroid and enlarged view of a polypite
showing pedicels ringed only at their bases.

ANTHOMEDfS.E — I'ENNAHI \. 27

water, and are often retained until they wither upon the stems after having cast out their
genital products.

I have found only the pale form of Pennarin on the Florida Reefs, and its medusae appear
to cast out their genital products before being set free, as is described by Hargitt for the deep-
water forms of Woods Hole, Massachusetts. In Florida, Pomona grows in tuft-like clusters
upon gorgonians. This tropical form is called Pennana gtbbosa by L. Agassiz, but those seen
by me can not be separated from the Pennaria of the coast of New England.

Hargitt, 1899, carried out a series of grafting experiments upon Pennaria and other
hydroids. He finds that pieces of hydroids of the same species may readily be grafted one
upon the other, both in oral or aboral relations, there being little or no evidence of polarity
in the regenerative process. Pieces of opposite sex but of the same species will readily unite
in any manner, but pieces belonging to hydroids of different genera would not unite. Elaborate
studies of this character were also carried out by Gast and Godewski, 1903, on P. Jisticha.

In 1900 and 1901, Hargitt studied the variations of the medusx and hydroids of Pen-
naria. The medusa sometimes displays ectodermal blister-like protrusions on its exumbrella,
and is variable in other respects.

According to H. Miiller, 1907 fZeit. fur wissen. Zool., Bd. 89), the eggs of Pennaria
develop at the expense of the weaker egg-cells of the ovary, which they engulf as food to form
the yolk-granules.

Thacher, 1903, shows that in Pennaria, Campaniilana, and EiiJenJnurn, the hydranths,
when they degenerate, are absorbed not by liquefaction of their protoplasm, or by the with-
drawal of the polyps as a whole; but absorption takes place by the degenerating cells of
both ectoderm and entoderm being turned into the digestive tract of the hydroid.

Goldfarb, 1906, finds that light is absolutely essential for the normal growth, develop-
ment, and regeneration of Pennaria. He finds, however, that this is true in a sense also for
Eudendrium, but not to the same degree, for Eudendrnim ramositm colonies, kept in the
dark until all the influence of their previous illumination has been lost, will not then
regenerate new hydranths unless they be again exposed to light; but the surprisingly short
exposure to light of only 5 seconds will suffice to restart the regenerative process.

Pennaria rosea von Lendenfeld.

Pennaria rosea, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, ser. I, vol. 9, p. 51)4, plate 24, figs. 40, 42. —
BALE, 1888, Ibitl., ser. 2, vol. 3, p. 747 — 7*. australii, BALE, 1884, in Cat. Australian Hydroid Zoophytes, p. 45.

Main axial stems 80 mm. long, with about 20 alternately arranged, pinnate side branches.
4 to 6 hydranths on each branch. Hydrorhiza and main stems intensely black and opaque;
outer half of each stem light-yellow, proximal half intense black. Hvdranths rose-colored,
with 9 to 14 oral, and 7 to 12 filiform, basal tentacles. Medusas are produced on the proximal
hydranths. The medusa bell is slender, oval, 2 mm. high, I mm. wide. 4 large rudimentary
tentacle-bulbs with a minute external ocellus upon each. Manubrium with sperm or ova fills
the entire cavity of subumbrella. Ova discharged after medusa is set free. Color, intense rose.

Coast of New South Wales, Australia. Mature in May.

This species is distinguished from the European and American Pennaria by its tentacu-
lar ocelli.

Pennaria adamsia von Lendenfeld.

Pennaria aaamsia, VON LENDENFF.LD, 1884, Proc. Linnean Soc. New South Wales, vnl. 9, p. 595, plate 25, figs. 45-48; plate 26,
fig- 49-

MeJusir. — The male medusae are 1.5 mm. long and only 0.7 mm. wide; while the female
medusas are 1.5 mm. long and as broad as they are high. The male medusas have 4 radially
situated marginal tentacles, about as long as the width of the bell. These tentacles have
well-developed basal bulbs with minute ocelli, and the tentacles move about freely. The
spermatozoa fill the space between the manubrium and the sides of the subumbrella, and are
discharged within an hour after the medusa has been set free. The female medusas are
broad, and the tentacles are mere rudiments without ocelli. The bell-cavity is filled with
ova, which are soon discharged.

28 MEDUS.E OF THE WORLD.

H ydroid. — The stems are 60 to 80 mm. long, zigzag, and with 15 to 20 alternate branches.
The longest branches are nearest the base and are about 12 mm. long. There are about 8
hydranths, 1.7 mm. long, on each branch. These hydranths are slender, the proximal ones
only about half as large as the distal primary one on each branch. There are 2 verticils of oral
tentacles, each with 4 tentacles alternating in position with those of the other row. These
oral tentacles are short and each ends in a nematocyst-knob. The uppermost row of 4 oral
tentacles is in the 4 perradii, and the lower row in the 4 interradii. There are 24 long
filiform tentacles in a single row at the broad base of the polypite. These are about as long
as the polypite itself. The perisarc of the hydrorhiza and main stems is opaque and black,
and the side branches are yellow. The hydranths are white.

Coast of New South Wales, Australia. Medusae are produced in March.

This species is distinguished by the minute ectodermal ocelli upon the tentacle-bulbs
of the male medusae, and by the length of its tentacles. There are also but 2 verticils of
oral tentacles.

Pennaria pacifica Clarke.

Pennaria pacifica, CLARKE, 1907, Mem. Museum Comp. Zool. Harvard Coll., vol. 35, No. l, p. 6, plate I, figs. 1-6.

Hydrocaulus 20 to 35 mm. high. Internodes of the main stem without annulations at
their distal ends, and with only I or 2 rings at their proximal ends. Branches alternate.
Peduncles taper slightly to the base where there are 2 or 3 annulations. Hydranths with
12 to 14 filiments, and about 16 capitate tentacles somewhat irregularly arranged.

Pinco Island, Gulf of Panama, tropical Pacific.

Distinguished only by its few annulations at the internodes of the main stem. It is
probably only a local variety of P. tiarella.

Pennaria ? vitrea Agassiz & Mayer.

Pennaria vitrea, AGASSIZ and MAYER, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161, plate I, figs, l, 2.

Bell is 3 mm. in height; walls thick and rigid. There are 4 rudimentary tentacle-bulbs.
Velum not prominent. The 4 radial-canals are straight and narrow. The manubrium in the
female specimen was retracted within the cavity of the bell, but it was flask-shaped in the
male and projected for a short distance beyond the velar opening. These conditions, how-
ever, may be merely different states of contraction and not constant differences; but as we
observed only two individuals, one a male and the other a female, we can not be certain upon
this point. The ova are large and pyriform, and are grouped in 4 interradially arranged
clusters within the manubrium. The mouth-opening is simple, and there are no fimbriations
or appendages.

Prominent circular muscles were observed in the ectoderm ot the cavity of the bell in the
female, but these were not seen in the case of the male. It is possible, however, that they
become apparent only during certain states of contraction. In the female the ova and tentac-
ular bulbs were flesh-colored, and the entoderm of the mouth was green. In the male the
tentacular bulbs were green, the entoderm of the manubrium pink, and the lips green.

Found among the Fiji Islands, off Kimbombo Island, November 25, and ofF Mbatiki
Island, December 5, 1897.

Not having seen the hydroid I am very doubtful concerning the generic identity of this
medusa with Pennaria.

Genus TRICHORHIZA Russell, 1906.

Trichorhiza, RUSSELL, 1906, Proc. Zool. Soc. London, p. 99, plate 5.

GENERIC CHARACTERS.

The medusa is similar to that of Pennaria, but with one tentacle-bulb somewhat better
developed than the 3 others. The hydranth is solitary, with a branched, filiform hydrorhiza.
The perisarc forms a cup into the cavity of which the hydranth may be partially retracted.
There are 2 verticils of tentacles, an oral and a basal; and the medusa-buds arise from the
sides of the hydranth between these 2 verticils. The type-species is Trichorhiza brunnea
Russell, from the Firth of Clyde, Scotland.

ANTHOMEDUS.E — TRICHORHIZA, STEENSTRUPIA.

29

Trichorhiza brunnea Russell.

Trichorhiza brunnta, RUSSKLL, 1906, Proc. Zool. Soc. London, p. 99, plate 5, figs. 1-2; Abstract, Pruc. Zool. Soc. London,
No. 26, p. 6, Feb. 13, 1906.

Mature medusa unknown. When newly set free from the hydroid the medusa is pyriform,
with a thin-walled bell, well-developed velum, and 4 rudimentary tentacle-bulbs, one of
which is larger than the others. The manubrium when extended is as long as the depth of
the bell-cavity, and it is cylindrical, narrow, and provided with a simple, circular mouth-
opening which is surrounded by nematocysts. There are 4 simple, straight radial-canals,
but no gonads. The manubrium and tentacle-bulbs are yellow, the former being faint and
the latter golden in color.

This medusa is liberated by a solitary hydroid which was found on June 29, 1905, at a
depth of 17 fathoms in Ethick Bay, Bute, Firth of Clyde, Scotland. The hydrorhiza of the

Trichorhiza was entwined among and
around the tentacles of Coryrnorpha
nutans. The hydroid is 1 1 mm. in total
length, the hydranth itself being only
1.5 mm. long. The hydrorhiza is long,
filiform, and sparingly branched, with
about 6 simple branches which arise
from its lower (aboral) half. The
coenosarc does not appear to extend
over this branched lower half of the
hydrorhiza. Above the upper end of
the hydrorhiza the perisarc extends to
form a cup-like hydrotheca, which
bears 4 transverse grooves. Immedi-
ately below this hydrotheca the peri-
sarc displays several longitudinal lines.
Hydranth conical, 1.5 mm. long and
0.8 mm. wide, with an oral circlet of 7
very short tentacles having knob-
shaped ends. There is also a basal
circlet of 12 filiform tentacles, which
are ringed with nematocysts and are
tapering, and fully as long as the hy-
dranth itself. 8 or 10 medusa-buds
arise at various levels from the sides
of the polypite between the basal and
oral circlets of tentacles.

The perisarc of the hydrorhiza is
straw-colored, and that ot the hydro-
theca chocolate-colored. Body of hy-
dranth pale reddish-brown, tentacles
translucent white.

This hydroid bears some resemblance to Blastothela of Verrill, 1878. Amer. Jour. Sci.,
ser. 3, vol. 1 6, p. 374.

FK.. 3. — Trichorhiza brunnea, after Russell, Proc. Zool. Soc., London, 1900.
Hydroid, and enlarged view of recently liberated medusa.

Genus STEENSTRUPIA Forbes, 1846.

Sleenslrufia, FOR BBS, 1846, Annals and Mag. Nat. Hist., vol. 1 8, p. 287.— LKI:CKART, 1856, Archiv. fiir Naturgesch., Bd.22, p. 29.
Slrrnstrupia + Euf>h\!a, HAF.CKEL, 1879, Syst. der Mcdusen, pp. 29, 3 i .— BROWN t, 1895, Trans. Liverpool Biol. Association,

vol. 9, pp. 247, 248.
Eu/ihysa ±Sticnitrufia, AI.ASSIZ, L., 1862, Com. Nat. Hist. I'. S., vol. 4, pp. 542, \^\. HAM KM , 1X64, Jena. Zeitsrli. fur

Naturw., Bd. I, pp. 338, 339. -FoRin.s, \X^, Hntisll N,iked-eved Medusa-, pp. 71, 72.
Euph\ia, A(.ASSIZ, A., 1865, North Amer. Acal., p. 1*0. UK. MOW. i'jc>4, Bull. Mus. Comp. Zool. at Harvard Coll., vol. VI.

p. 251. — VANHOKFFN, 1891, Zool. An/eiger, lid. 14, p. 443.
Ku/>h\sora, MAAS, 1905, Hydromeduscn der Siboga Kipediiion, p. 6.
Heterosirf>hanus + Stcenslrupia+ Eaf>h\sa, HARTLALB, 1907, Nordischcs Plankton, Nr. 12, pp. 74, 76, 80.

30

MEDUS.E OF THE WORLD.

The type-species ot this genus is Steenftrupia rubra Forbes, 1846, from the Atlantic
coasts of Europe.

GENERIC CHARACTERS.

Codonids with radially symmetrical bell, and with 4 radially situated tentacles, I of
which is well developed while the 3 others are short or rudimentary.

This genus is separated from Hybocodon by its radially symmetrical bell. In Hybocodon
i radial-canal is long, I short, and 2 of medium length; but in Steenstrupia the 4 radial -canals
are all of the same length.

Synopsis of the Characters of MeJuste of the Genus Steenstrupia.

S. rubra Forbes=

S. aurata= Euphy-

S. tetrabrachia=E.

S. bigelowi Maas.

S. australis von

S.flaveola Forbes

sa aurata Forbes,

tetrabrachiaH.B.

Lendenfeld.

= S. galanthus

= E. aurata+E.

Bigelow.

Haeckel =(?)£.

mediterranea

gracilis Brooks=

Haeckel = (?) E.

( ?) S. cranoides

virgulata A.

Haeckel=S. 1m-

Agassiz.

eata Leuckart.

Shape and size

Bell has well-devel-

Dome-like apex.

Pyriform. 4 high,

Pyriform. 10 high,

Half-egg-shaped,

of bell in mm.

oped apical pro-

Cylindrical sides.

2.5 wide.

5 wide.

2.5 high, 1.7

jection. 5 high,

6 high, 4.5 wide.

wide.

3 wide.

Bell-walls and

apex thick.

Condition of ten-

3 long, narrow ten-

3 small, similar,

3 short tentacles,

3 short tentacles

3 tentacle-bulbs,

tacles. Length

tacle-bulbs with-

rudimentary ten-

each= r long, i

about 2 r long

and i long tenta-

in terms of bell-

out tentacles, all

tacle-bulbs, i

large tentacle

and not ringed.

cle, ringed with

radius (r).

similar, i long

long tentacle.

about 7 T long.

One longtentacle

nematocysts.

tentacle ringed

Length = 2 r+ .

All tentacles

4 T long, with

Long tentacle

with nematocysts.

No ocelli. No

sparsely ringed.

many prominent

about 8 r long.

Length = 7 r.

definite rings of

partial rings of

nematocysts on

nematocysts.

tentacles.

Axial-canal above

Axial canal always

No axial-canal.

No axial-canal.

Present. Small,

No axial-canal.

stomach.

present.

slender.

Length of manu-

1 .5 r. Stomach

1.25 r. No ped-

2. 5 r long. Mounted

1.5 r long. Mouth

1.5 T long. Cylin-

brium in terms

mounted on short

uncle.

on very short ped-

not quite reach-

drical.

of bell-radius

peduncle.

uncle ? Mouth

ing level of velar

(<•).

extends slightly

opening.

beyond velar

opening.

Gonads.

Single gonad en-

Single gonad en-

Eggs developed in 4 wide interradial,

Gonad encircles

circles stomach,

circles stomach.

8 adradial, longi-

ectodermal swell-

manubrium.

leaving mouth and

tudinal rows, in

ings on sides of

peduncle free.

ectoderm on sides

mid-region of man-

of stomach.

ubrium. Separated

by 4 narrow, per-

radial spaces.

Color.

Tentacle-bulbs and

Variable. Tenta-

Gonads brownish-

Entoderm of man-

Mouth deep violet.

stomach light-

cle-bulbs may be

yellow. Mouth

ubrium, canals,

4 brown patches

pink to reddish-

yellow, orange,

pink. Tentacle-

and tentacles dull

on gonad. Tenta-

brown. Bell apex

red, or colorless.

bulbs and rings

yellow .

cle-bulbs and long

sometimes faint

Manubrium red-

pink.

tentacle brown

pink .

dish to yellow or

with violet spots.

faint purple.

Where found.

Atlantic coasts of

Mediterranean and

Suvadiva Atoll,

Malay Archipelago.

Harbor of Sydney,

Norway, Ger-

Atlantic coasts of

Maldive Islands,

New South

many, England,

Europe.

Indian Ocean in

Wales, Australia.

Scotland, and Ire-

January.

land. (Mediter-

ranean?) (South-

ern United States,

Atlantic coast?)

Hydroid.

Corymorpha nu-

Corymorpha nana,

Unknown.

This medusa is pro-

Unknown.

tans, Sars, 1835;

Alder, 1857.

bably identical

Hincks, 1868;

with S. tetrabra-

Allman, 1871.

chia. Hydroid

unknown.

ANTHOMEDUS.E — STEENSTRUPIA. 31

The hydroid generation is Corymorpha Sars, i$3$ = Halatractus of Allman, 1871.

Haeckel, 1879, restricts Steenstrupia to describe medusae having characters as above,
but with an apical projection upon the bell, and with an axial canal extending upward into
this projection from the stomach. Euphysa he would restrict to include forms which lack an
apical projection and an axial canal. As Vanhoffen, 1891, rightly states, an apical projection
is always lacking in young medusae and is highly variable in its development even in mature
individuals of the same species. The same is true of an axial canal. Moreover, among
species discovered since Haeckel wrote his "System der Medusen," Euphysa tetrtibrachin,
Bigelow, 1904 (Bull. Mus. Comp. Zool. at Harvard Coll., vol. 39, p. 251, plate I, fig. i), has
a well-developed apical projection, and no axial canal, and would therefore have no place in
Haeckel's system.

Maas, 1905, would institute a genus Euphysora to describe Codonidae having I large,
and 3 well-developed but still considerably smaller tentacles. Here again, however, we meet
with a condition of degree only. When, for example, are the 3 small tentacles large enough
to be Euphysara or small enough to be Steenstrupia ? In order to avoid confusion, it appears
best to combine all of these forms under one generic name. Vanhb'ffen, 1891, suggests Euphysa
as the proper generic name to include all of these forms, but Steenstrupia takes precedence
over Euphysa, for it was used by Forbes in 1846, while Euphysa was first used by him in 1848.

Bigelow finds that the eggs of S. tetrabrachia are arranged in 8 adradial longitudinal lines
in the ectoderm of the stomach, and Maas states that in S. bigelou'i the gonads are interradial
and separated by very narrow linear interspaces in the 4 principal radii. These conditions
are interesting, for they foreshadow those characteristic of the Oceanidae where the gonads
are interradial or adradial, and are often quite widely separated in the principal radii.

Hartlaub, 1907, defines Heterostephanus (Allman, 1871) as a Steenstruf>ia-\\ke medusa-bud,
not known to be set free from its Coryrnorpha-\\\ie hydroid. Medusa with a single tentacle,
ringed at its base, and terminating distally in a knob. The only known species is H . annuli-
cornis, Allman, l^l=Hybocodon annulicornis Haeckel. This was first described by M.
Sars, 1859; as Corymorpha ? annulicornis.

Steenstrupia rubra Forbes.

Plate I, fig. 7.

Corymorpha nutans (hvdroid), SARS, M., 1835, Beskriv. og Jagtt., p. 6, plate i, fig. 3; SARS, O., 1877, Fauna Littor. Norveg.,

tome 3, p. 2, taf. 2, figs. 25-28.

Corymorpha nutans (hvdroid), BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 63 (literature to 1850).
Corymorpha nutans (hydroid and medusa), HARTLATB, 1907, Nordisches Plankton, Nr. 12, p. 76, figs. 72-75 (complete li

authors, localities, and best modern description of hydroid and medusa).
Corymorpha nutans, HINCKS, 1868, Hist. British Hvdroid Zoophytes, p. 127, plate 22. — AI.LMAN, 1871, Monog. Tubularian

Hydroids, p. 388, plate 19.
Corymorpha nuians = S. galanthus (Haeckel)=.S. faveola, Forbes = S. rubra, Forbes, BROWNE, 1896, Proc. Zool. Soc. London,

pp. 463, 464, plate 16, fig. i (review of literature).
Steenstrupia galanthus, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 148 (literature to 1850). — HAECKEL, 1879, Syst. ilrr

Medusen, p. 31. Hybocodon ntttans, Ibid., p. 34. — MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, p. 52, taf. 5, figs. 20-24

(origin and structure of the eggs).
Steenstrupia rubra, HARTLAUB, 1904, Wissen. Meeresuntersuch. Kommiss. Meerc Kiel, Abth. Helgoland, Neue Folgc, Bd. 5,

p. 105, fig. 3.

Steenstrupia rubra (Forbes), BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, p. 247.

Steenstrupia rubra (medusa)+S. faveola, FORBES, 1848, British Naked-eyed Medusa, pp. 73, 74, plate 13, figs, i, 2.
Steenstrupia cranoides, HAECKEL, 1879, Syst. dcr Medusen, p. 30, taf. 2, figs. 10-14.
Steenstrupia gracilts, BROOKS, 1882, Studies Johns Hopkins Vniv. Biul. Laboratory, vol. 2, p. 144. — MAYER, 1900, Bull. Mus.

Comp. Zoo!, at Harvard Coll., vol. 37, p. 29, plate 16, figs. 36, 37.
Steenstrupia lintata, LEUCKART, 1856, Archiv. fur Naturgesch., Bd. 22, p. 29, taf. 2, fig. 6. — SPACNOLINI, 1876, Catali-i:" \

Mediterraneo, p. 17, taf. I, figs. 1-4. — HAECKEL, 1879, Syst. der Medusen, p. 30. — DU PLESSIS, 1888, Recueil /.<«.!. s,n re,

tome 4, p. 543.

This medusa is found on the coasts of Europe from Norway southward to the Mediter-
ranean. S. cranoides Haeckel = .V. lineata Leuckart, from the Mediterranean, appears to be
identical with S. rubra. S. gracilis Brooks, of the Atlantic coast of the United States south
of Virginia, is probably identical with S. rubra of Europe, but the hydroid of the American
form remains unknown. I believe that 5. Un<-ntn, S. rrtinoiJfs, and .V. i>r/ii-ilis are identical, and
that they are probably identical with .V. ruhrn Forbes.

For description of i'. rubra of Europe, see synoptic table of species of Steenstrupia.
3

32

MEDUSAE OF THE WORLD.

In the American form the development of nematocyst-rings upon the long tentacle is
subject to great variability. The Mediterranean S. cranoides Haeckel lacks such rings,
whereas they appear to be better developed in northern Atlantic specimens from the coasts
of Europe. The same difference appears to be exhibited by our American specimens, those
from Tortugas, Florida, being unnnged or only slightly ringed, while those from North Car-
olina are often profusely ringed. The apex of the bell often bristles with nematocysts, but
in some medusae it may be smooth.

Browne, 1896 (pi. 16, fig. i), gives a figure of S. rubra derived from specimens found by
him at Valencia Island, off the Irish coast. He shows a narrow conical peduncle above the
stomach, the peduncle being about one-fifth as long as the gastric portion of the manubrium.
No such peduncle has been observed in the American S. " gracilis," when the bell is expanded,
although when it is somewhat contracted the appearance of a well-developed peduncle is
often produced. On the other hand, when the stomach is widely distended with food and the
bell expanded no peduncle may be seen. Haeckel's series of figures (taf. 2, figs. 10-12) will

3>

FIG. 4. — Corymorpha nulans, after Hartlaub, in Nordisches Plankton.
(Young hydroids and mature medusa.)

serve to illustrate the formation of a peduncle-like body of gelatinous substance above the
stomach through contraction of the bell. I have frequently seen the same phenomenon in
our American S. " gracilis" = S. cranoides Haeckel. I have also observed this peduncle in
living medusae of S. rubra taken off the coast of Cornwall, England.

Hartlaub, 1907, gives a list of the bibliography and of localities for this species, and his
description of the medusa and the young hydroid are the best yet produced (see fig. 4).

The egg is amoeboid as in Amalth&a. The young hydroid has a single circlet of 4 short,
knobbed, oral tentacles, and another circlet of 5 to 8 simple, flexible, filiform basal tentacles.
H. Miiller, 1908, finds that the full-grown eggs are very few in number, having developed at
the expense of other weaker egg-cells in the ovary, which they devour. The exoplasma is quite
wide and is separated from the germinal vesicle. The ooplasma is a network of delicate fibers
of wide mesh, and the exoplasma and endoplasma are distinct, one from another. The egg
contains numerous pseudo-cells in advanced stages of degeneration and also yolk-granules.

ANTHOMEDUS.E STEENSTRUPIA.

33

The following description of the medusa is derived from a study of specimens found by
the author off the coast of the United States:

The bell is 5 mm. high and surmounted by a slender conical projection about 2 mm.
long. There are 4 tentacles. One of these is about 10 mm. long, and is ringed at irregular
intervals by prominent swellings, between which there are small rings at fairly regular inter-

FIG. 5. — Corymorfha nuians, hydroid and medusa, after Allman, in Ray Society, 1871-72.
A' Detail showing manner in which medusa1 bud off from hydranlh.

34

MEDUSAE OF THE WORLD.

vals. The tentacle which is diametrically opposite to the long tentacle is tapering, and about
0.25 mm. long; while the 2 other tentacles are mere bulbs. The velum is well developed.
There are 4 narrow radial-canals, and a slender ring-canal. A long, slender, sinuous
canal extends from the aboral apex of the stomach upward into the apical projection of the
bell. In mature medusae the manubrium extends a short distance beyond the velar opening.
Ordinarily the mouth is a simple, round opening at the tapering extremity of the manubrium,
but when widely open, as in our figure, it shows 4 thick but not prominent lips. The genital
products are found in the manubrium, and in the female the eggs project from the surface of
the ectoderm. Entoderm of manubrium intense yellow-green and rose-color. Apical canal
intense yellow, often flecked with rose-color. The entoderm of the tentacle-bases is rose-
color and yellow, while the entoderm of the large annular swellings is rose-color. When
young the apical projection is not very high, the tentacles short, and without nematocyst-
nngs, and the manubrium short and tapering; not extending beyond the velar opening as
in the mature medusa.

6.

FIGS. 6 and 7. — Sieenslrufia rubra.

6. — From life, by the author. Off Mousehole, Cornwall, England, October 23, 1907.
7. — A. After Leuckart ("S. lineata"), 1856, Archiv. fur Naturgesch., Bd. 22.
B. After Spagnolini ("S. lineata"*), 1876, Catalogo Acalefi Mediterraneo.

Found at Oregon Inlet, Pamlico Sound, North Carolina, in November, and at Beau-
fort, North Carolina, and Tortugas, Florida, in summer. It is apparently identical with
S. cranoides and S. lineata of the Mediterranean.

Definite rings of nematocysts are not found upon the tentacles of the young medusa, and
are very variable in their development in mature specimens, some being profusely ringed
and some entirely unringed. Haeckel describes only unringed individuals from the Medi-
terranean, his specimens being similar to those found by me at Tortugas, Florida.

I am inclined to believe that this Mediterranean and tropical American medusa will
prove to be identical with, or only a variety of, S. rubra of the Atlantic coasts of western
Europe. Certainly no differences, other than those well within the limits of common vari-
ability, can be detected between the medusae of S. rubra and S. lineata = S. cranoides; but a

ANTHOMEDUS.E — STEENSTRUPIA. 35

careful comparative study of the hydroids of these forms must be made before we may safely
assert either that they are identical or separate species.

The hydroid of S. rubra is Corymorpha nutans of Sars, and is common on sandy bottoms,
off the northern coasts of Europe, at moderate depths. The stems of the hydroid are
solitary, and are about 50 to 75 mm. high, and about 4 mm. wide at the widest part. The
whole stem is corrugated by numerous narrow longitudinal bands. The widest part of the
stem is usually at a short distance above the lower end. This lower end is bent sharply at
right angles to the main part of the stem and bears long, hair-like filaments which serve to
anchor the hydroid. There are also blunt, papilla-like processes which arise from the sides
of the stem above the bent portion. The polypite is large and flask-shaped, and has a basal
zone of 30 or more long, tapering tentacles, about as long as the polypite itself. In addi-
tion to these tentacles there are 6 to 7 closely crowded verticils of oral tentacles, which are
much shorter and thinner than the proximal. 15 to 20 branched peduncles arise from the
sides of the polypite close to the bases of the proximal circlet of tentacles, and bear the
medusa-buds. The hydranth is light-red, the stem being paler than the polypite. On the
English coast the medusa-buds are set free during the summer.

Allman gives a good description of the hydroid.

Steenstrupia aurata.

Euphysa aurata, (medusa) FORBES, 1848, British Naked-eyed Meduss, p. 71, plate 13, fig. 3.

Euphysa aurata (medusa)+E. mcditcrranea, HAECKEL, 1879, Syst. der Medusen, p. 32, taf. 2, figs. 8, 9.

Euphysa medilrrranca, DU PLESSIS, 1888, Recueil Zool. Suisse, tome 4, p. 543.

Euphvsa aurata (Forbes), BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, p. 248; 1896, Proc. Zool. Soc. London, p. 474.

Euphvsa aurata, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 749. — BEDOT, 1905, Revue Suisse de Zool., tome 13,

p. 134.
Corymorpha nana, (hydroid) ALDER, 1857, Cat. Zooph. Northumberland and Durham, p. ill, plate 7, figs. 7, 8. — HINCKS, 1868,

Hist. British Hydroid Zooph., p. 130, plate 22, fig. 3.

Corymorpha nana, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 81, figs. 76-78 (list of authors and localities).
(?) Euphysa virgulata, AGASSIZ, A., 1865, North Amer. Acal., p. 189, figs. 316-319.

This medusa is found off" the Atlantic coasts of western Europe and in the Mediter-
ranean. Steenstrupia virgulata of Massachusetts Bay is probably identical with S. aurata.
For description of the European form, see synoptic table of characters of the species of Steen-
strupia. The European S. aurata appears to be smaller than the American S. virgulata.

Steenstrupia virgulata = (?) S. aurata Forbes.
Plate i, fig. 6.

(?) Steenstrupia aurata, FORBES, 1848, British Naked-eyed Medusae, p. 71, plate 13, fig. 3.

Euphysa virgulata, AGASSIZ, A., 1865, North Amer. Acal., p. 189, figs. 316-319. — HAECKEL, 1879, Syst. der Medusen, p. 33. —
NUTTING, 1901, Bull. U.S. Fish Com m., vol. 19^.370. — HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 33.
Corymorpha virgu/aia, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 84, fig. 79.

Adult medusa. — The bell is pyriform, with a broad, dome-shaped apex. It is 5 to 12 mm.
in height, and 4.5 to 9 mm. in diameter. Surface of exumbrella smooth and without rows of
nematocysts. There are 4 tentacles, one at the base of each radial-canal; 3 of these are mere
rudimentary bulbs, but the fourth is large and conical. Its surface is thickly covered with
nematocyst-cells of large size. There are 4 simple, straight radial-canals, and a slender,
circular vessel. The velum is wide, with an indented, free edge. The manuhrium is cylin-
drical without a peduncle, and extends about half the distance from the inner apex of the
bell-cavity to the level of the velar opening. No apical, axial canal. The mouth is a
simple, round opening without prominent lips. The genital products are contained within
the ectoderm of the manubrium. Manubrium light-yellow. The entoderm of the radial-
canals near the bases of the tentacles is intense pink, and the ectoderm of the tentacles is
milky-white. This species was found by Dr. Alexander Agassiz, in Massachusetts Bay, at
Nahant, and is recorded from Woods Hole, Massachusetts, by Nutting and Har^itt.

The figure here shown is reproduced by his kind permission from Dr. Alexander Agassiz's
drawing of the medusa. There is no difference between this medusa and S. aurata of Europe
except that the American form appears to be larger; but the hydroid of the American medusa
is unknown, and until this is discovered it will be impossible to determine the identity of the
American form.

36

MEDUSAE OF THE WORLD.

Steenstrupia tetrabrachia.

Euphysa telrabrachia, BIGELOW, H. B., 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251, plate I, fig. I.
(?) Euphysora bigelowi, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 7, taf. I, figs. 1-3.

Bell 4mm. high, 2. 5 mm. wide. Pyriform,with solid apical projection. One long sparsely-
ringed tentacle about 4 times as long as the bell is high. 3 other, smaller tentacles, each about
one-third as long as the bell-height. These bear each about 3 rings of nematocysts, while the
long tentacle bears about 6 such rings. Velum well developed. Manubrium large, spindle-
shaped, with mouth projecting beyond the velar opening. Gonads on sides of stomach.
Eggs arranged in 8 fairly distinct rows. Bell colorless. Gonads brownish-yellow. Manubrium
pinkish. Tentacle-bulbs and rings on tentacles rose-pink. Suvadiva Atoll, Maldive Islands,
Indian Ocean; in January.

This medusa is probably redescribed by Maas as E. bigelowi.

Steenstrupia bigelowi.

Euphysora bigelowi, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 7, taf. i, figs. 1-3; 1906, Revue

Suisse de Zool., tome 14, p. 84, pi. 2, figs, i, 2. — MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, p. 59.
(?)Euphysora tetrabrachia, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251, plate i, fig. I.

Fir,. 8. — Steenstrupia tetrabrachia, after Bigelow,
in Bull. Museum Comp. Zool. at Harvard
College.

FIG. 9. — Steenstrupia bigelowi, after Maas, in
Hydromedusen Siboga Expedition.

Bell 13 mm. high; more than twice as high as it is wide, and with a well-developed
apical projection. Bell-cavity 10 mm. deep. Apical projection 3 mm. high. Side walls thin.
4 tentacles at the bases of the 4 radial-canals. One of these tentacles is longer than the bell-
height and bears about 30 swollen nematocyst-warts. The 3 other tentacles are each one-
third to half as high as the bell and are tapering, without nematocyst-warts, but covered with
diffuse nettling cells. Manubrium spindle-shaped, not extending beyond the velar opening.

ANTHOMEDUS.E — STEENSTIU'I'IA, HYBOCODON. 37

In most specimens there is a blindly-ending axial canal extending into the gelatinous substance
of the apical projection, but this is not constantly present. The gonads are developed upon
the sides of the stomach, leaving only the basal and mouth ends of the manubrium free. A
cross-section shows that they are separated by 4 minute, perradial, longitudinal lines. There
are therefore 4 interradial gonads. There are no medusa-buds produced by the medusa.

The entoderm of the manubrium, radial-canals, and tentacle-bulbs is filled with yellow
pigment granules. The nematocyst clusters on all 4 tentacles are red.

Found quite commonly in the Malay Archipelago, at Sulu, Ternate, Damar, Manifa,
Saleyer, and Amboina.

This form differs from the closely allied S. letrabrachia Bigelow, from the Maldive
Islands, in that in the Maldive species there are a few rings of nematocysts upon the ten-
tacles, whereas in S. bigelou'i there are only warts, not inclosed rings. Also there appears to
be no axial canal in S. tetrabrachia, whereas this is usually seen in 5. bigelowt. The size
constitutes a disparity in the two medusae 4 mm. high in S. tetrabrachia and about 13 mm. in
S. bi^elowi. Future studies will probably show that these distinctions are not of specific
value, but merely changes due to growth and variation, and that the two medusae are identical
and should be called S. tetrabrachia. "Euphysa" tentaculata, Linko, 1905 (Zool. Anzeiger,
Bd. 28, p. 214), from Barents Sea has also 3 well-developed tentacles, and is 5 mm. high,
with orange-colored manubrium and oval bell. It may be Hybocodon pcndula.

Steenstrupia australis.
Euphysa ausiralis, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 586, plate 21, fig. 33.

Bell is 2.5 mm. high, 1.7 mm. wide. Half-egg-shaped and symmetrical. No lines of
nematocysts over the exumbrella. One very long retractile tentacle, 2 to 3 times as long as
the bell-height. This tentacle has a large basal bulb, and is covered with rings of nemato-
cysts. The other 3 tentacles are mere basal bulbs terminating in a knob-shaped cluster of
nematocysts. Velum well developed. 4 straight radial-canals. Manubrium arises from the
center of the umbrella cavity, and is cylindrical and about half as long as the bell-height.
The gonad encircles the manubrium. Mouth deep violet. 4 brown patches upon the gonad,
and a few brown spots on manubrium near its base. Tentacle-bulbs and the large tentacle
brown with violet spots. Port Jackson, New South Wales, in May and June. Rare.

Hydroid unknown.

Genus HYBOCODON L. Agassir, 1862.

Htbocodon, AGASSIZ, L., i86z, Cont. Nat. Hist. U. S., vol. 4, p. 243.— AGASSIZ, A., 1 865, North Amcr. Acal., p. 193.— VANHOFFEN,
1891, Zool. Anzeiger, Bd. 14, p. 443. — BROWNE, 1896, Proc. Zool. Soc. London, p. 466. — HARTLAI B, 1905, Zoolog. Jahr-
buchern, Suppl. 6, p. 544; 1907, Nordisches Plankton, Nr. 12, p. 96.

HybocoJon + ArnphicoJon, HAECKF.L, 1879, Syst. der Medusen, pp. 33, 35.

Amphirodon, BROWNE, 1901, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 275.

Diplura, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 326.

Corymorpha, AGASSIZ, A., 1865, /feiW., p. 192.

This genus was established in 1862 by L. Agassiz for Hybocodon prohfer, a medusa
which arises by budding from a Corymor^/ja-like hydroid on the New England coast. Accord-
ing to Browne, 1896, this medusa is also found off the northern coast of Europe. It is prob-
able that the same medusa was described by Steenstrup, 1842, from Iceland, as Corymorpha
fritillaria, but the hydroid from which Steenstrup supposed this medusa to be derived is
certainly not HvbocoJon, but may be an Arnalthira or Diplura. Steenstrup does not figure
a basal circlet of tentacles upon the polypites, which have only an oral circlet, and below
this a circlet of medusiform gonophores, each with a 4-sided bell, and 4 equally developed
rudimentary tentacles. It is probable that the hydroids of Hybocodon differ more among
themselves than do the medusae, and no final classification of the medusae can be attempted
until all of the hydroids have been discovered.

GENERIC CHARACTERS.

Codonidae with asymmetrical bell. One of the 4 radial-canals is long. I short, and 2 of
medium length. There are I or more long tentacles at the foot of the long radial-canal;
and 3 small or rudimentary tentacles, I at the foot of each of the 3 other radial-canals. The
hydroid is HybocoJon.

38

MEDUS.E OF THE WORLD.

Haeckel, 1874, instituted the genus Amphicodon to include medusae with 3 rudimentary
tentacles, and a cluster of 2 or more long tentacles at the foot of the longest radial-canal.
Vanhoffen, 1891, Browne, 1896, and Hargitt, 1901, have pointed out, however, that the
young medusae commonly have but I long tentacle, and that others appear, and develop from
the side of the basal bulbs of this original tentacle; thus the genus "Amphicodon" is only
a mature Hybocodon. These secondary tentacles may appear before or after the medusa-
buds begin to develop upon the tentacle-bulbs. According to Hargitt, 1902, 1904, Perkins,
1904, and Linko, 1905, the sexual products of the manubrium become mature while
medusa-buds are still being produced upon the tentacle-bulbs.

Synopsis of the Species of Hybocodon.*

H. prolifer L. Agassiz.

H. pendula Haeckel =
Corymorpha pendula
L. Agassiz.

H. forbesii Mayer.

H. unicus Browne.

Shape and size of bell

Dome-shaped. 2.5 high,

Quite similar to H. pro-

Ellipsoidal, asymmetri-

Bell-shaped. 3 high,

in mm.

2.2 wide, i radial-

lifer. 5 high, 3.5

cal. 2.5 high, 2.1

2 wide.

canal long, 2 inter-

wide.

wide.

mediate length, and i

short. Long and short

are 180° apart.

Number of longitudi-

2 extend upward from

As in H. prolifer.

None.

?

nal lines of nemato-

base of well-developed

cysts on exumbrella

tentacle, and I from

base of each of 3 rudi-

mentary tentacle-bulbs.

Condition of well-de-

i to 3 well-developed ten-

Only i well-developed

Only i well-developed

Only i well-developed

veloped tentacle, or

tacles at base of longest

tentacle. No medusa-

tentacle. No medusa-

tentacle "between 2

cluster of tentacles,

radial-canal. Medusa-

buds.

buds.

rudimentary basal

at base of longest

buds are produced at

bulbs." No medusa-

radial-canal.

bases of these tentacles.

buds.

Often only i well-

developed tentacle pro-

duced.

Condition of 3 rudi-

Mere basal bulbs.

i mere basal bulb. 2

i short conical tentacle.

3 mere basal bulbs.

mentary tentacles.

small tentacles. Small

2 mere basal bulbs.

tentacles 90° and

Short conical tentacle

basal bulb 180° from

1 80° and basal bulbs

large tentacle.

90° from long tentacle.

Gonads.

Developed over ectoderm

On ectodermal sides of

On manubrium. No

?

of manubrium. Actinula

manubnuni. No ac-

actinulx attached to

larvs develop upon sur-

tmulie seen attached

gonad.

face of gonad in ecto-

to gonad.

derm of manubrium.

Color.

Entoderm of tentacle-

Pink granules in tenta-

Entoderm of tentacles

?

bulbs, radial-canals,

cle-bulbs. Entoderm

and manubrium yel-

and stomach orange to

of manubrium pink

low, flecked with red.

deep blood-red.

and lilac.

Where found.

North Atlantic coasts of

Coast of New England,

Bahama Islands, and

Stanley Harbor, Falk-

Europe, Iceland, and

United States.

Florida coast. Com-

land Islands.

America.

mon in spring months,

at surface.

Hydroid .

Hydroid described by L.

Hydroid is Hybocodon

Unknown.

Hydroid may be H.

Agassiz, 1862, as H.

pendula= Corymorpha

chilensis Hartlaub,

prolifer.

pendula L. Agassiz.

1905.

*For description of H. chilensis Hartlaub, H. chrislinf Hartlaub, H. pu/cher Hartlaub, and H. (?) januarii, see text.

Hybocodon prolifer L. Agassiz.
Plate 2, fig. i; plate 3, fig. 3.

(?) Corymorpha fritillaria, STEF.NSTRUP, 1842, Generations-wecksel., p. 20, taf. I, figs. 41-46 (the medusa only may be identical

with //. prolifer; the hydroid appears to be an Amaltheea or Diplura).
( ? ) Stcenstrupia globosa, SARS, 1859, Christiania Vidensk. Selsk. Forhandl., p. 101. — KOREN and DAWELSSEN, 1877, in Sars's

Fauna Littoralis Novegiie, tome 3, p. 20, taf. i, figs. 1-6.
(?) Corymorpha annulicornis (young medusa), KOREN and DANIELSSEN, 1877, Ibid., p. 8, taf. i, figs. 7-13.

PLATE 2.

Fig. I. Hybocodon prolifer. Woods Hole, Massachusetts, March 4, 1907.
This drawing shows one of the normal aspects of the medusa,
with the bell somewhat contracted. When the bell is extended the
mouth comes to the level of the velar opening.

Fig. 2. Hybocodon pendula. Newport, Rhode Island, April 23, 1897.

Fig. 3. Hybocodon forbesii. Tortugas, Florida, June 26, 1906.

Fig. 4. Dicodoniurn jeffersoni. Tortugas, Florida.

Fig. 5. Dicodoniurn floriJana. Tortugas, Florida, June 17, 1897.

Drawn from life, by the author.

PLATE 2

ANTHOMEDUSJE — HYBOCODON. 39

Hybocodon prolijer (hydroid and medusa), AOASSIZ, L., 1861, Cent. Nat. Hist. U. S., vol. 4, pp. 143, 343, plate 133, figs. 10,

11; plate 25, 19 figs.
Hybocodon proltffr, AGASSIZ, A., 1865, North Amer. Acal., p. 193, figs. 325-328. — ALLMAN, 1871, Monog. Tubul. Hydroids,

p. 422. — VERRILL, 1873, Invert. Animals, Vineyard Sound, p. 736, plate 38, fig. 282. — BOHM, 1878, Jena. Xritschr. fiir

Naturw., Bd. 12, p. 195, taf . 7j figs. 7-9. — NUTTING, 1901, Bull. U.S. Fish Commission, vol. 19, p. 341, fig. 76. — HARGITT,

1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 33, plate 2, fig. 2; 1901, American Naturalist, vol. 35, p. 580, fig. 39;

Ibid., 1902, vol. 36, p. 552; 1901, Biol. Bulletin Woods Hole, vol. 2, p. 222. — PERKINS, 1904, American Naturalist 38,

p. 516 (simultaneous sexual and asexual reproduction).
(?) Hybocodon prolijer, BROWNE, 1896, Proc. Zool. Soc. London, p. 466.
Hybocodon prolifer + H.annulicornis + ? Amphicodon fritillaria+ ? A. giobosus + A.amphipleurus, HAKCKEL, 1879, Syst. JerMcdu-

sen, pp. 33, 35, 36, 37, taf. I, figs. 7-9.

(?) Amphicodon fntillaria, BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9.

(?) Amphicodon grtn°idum, LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 215 (simultaneous sexual and asexual reproduction).
(?) Hybocodon prolifer, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 752.
Non Hybocodon prolijer, BONNEVIE, 1899, The Norwegian North Atlantic Expedition, 1876-1878, vol. 26, Hvdroida, p. 28, plate

i, fig. 6.
Hybocodon prolijer, HARTLAUB, 1907, Nordisches Plankton, Mr. 12, p. 98, figs. 94-97. — MULLER, H., 1908, Zeit. fur wissen.

Zool., Bd. 89, pp. 62, 73 (origin and structure of the eggs).
( ?) Hybocodon gravidum + H. islandicus + H . amphipleurus, HARTLAVB, 1907, Nordisches Plankton, Nr. 12, pp. 104, 106,

figs. 99, 100.

The following description is derived from studies made of medusae from the southern
coast of New England, United States:

Adult medusa. — Bell about 2.5 mm. high and 2.2 mm. wide. It is asymmetrical, the side
bearing long tentacles being longer than the other sides; or as Browne aptly describes it,
the margin is not at right angles to the longitudinal axis of the bell, but slopes toward the side
bearing the cluster of long tentacles. The 4 radial-canals are of lengths corresponding to
the sides of the bell. The canal leading to the cluster of long tentacles is the longest; while
the canal diametrically opposite to this is the shortest, the 2 other canals being of inter-
mediate length. There are 3 small, rudimentary tentacle-bulbs, I at the foot of the shortest
canal, and I at the foot of each of the intermediate canals. The cluster of tentacles at the
foot of the long radial-canal has wide, hollow, tapering basal bulbs. The mam shaft of each
tentacle is, however, slender, cylindrical, and contractile, and is annulated at regular intervals
by well-developed clusters of nematocysts. Young medusae commonly have but a single long
tentacle, but as growth proceeds they sometimes develop another and finally a third; and
all 3 grow to be of equal length, and form a conspicuous cluster. A number of medusa-buds
in various stages of development arise from the sides of the hollow base of the one or more
long tentacles, near the bell-margin. These medusa-buds themselves develop a single long
asymmetrical tentacle even before the bud is mature. When ready to be set free they resemble
the parent medusa in that they are sometimes observed to be developing a third generation
of medusae upon their tentacle-bulbs. 5 longitudinal lines of nematocyst-cells extend from
the bases of the tentacles to near the apex of the bell. 3 of these rows arise from the bases
of the 3 rudimentary tentacles, and extend up over the surface of the exumbrella immedi-
ately over the radial-canals. In addition to these there are 2 rows which start from the base
of the well-developed tentacles, and extend upward over the exumbrella surface on both
sides of the long radial-canal. The velum is well developed. The radial-canals are narrow
and straight. The manubrium is a simple tube, which usually extends about two-thirds
the distance from the inner apex of the bell-cavity to the velar opening. The mouth has
4 short lips with their edges surrounded by nematocysts. The mature eggs are found in the
ectoderm of the stomach, and there they develop into actinula larvie before being set free.
The entoderm of the tentacle-bulbs is intense orange. The rows of nematocysts upon the
exumbrella often display an orange tinge, as does also the entoderm of the stomach.

Hydroid. — The hydroid was found in Massachusetts Bay. growing in tide-pools where
the water was very pure. Stems about 50 mm. in height. They grow singly, or in small clusters,
and do not branch. Each stem terminates distally in a single large polypite. The stems
are not more than I mm. in diameter at the base, gradually enlarging toward upper end,
and are about 3 mm. in diameter at base of polypite. They are covered with a delicate sheath
of chitinous perisarc, which widens and displays several well-developed annulations near
the base of the polypite. The polypite is flask-shaped with a very broad base; the mouth is
situated at the extremity of a narrow cylindrical neck, which is capable of much distension.
There are 2 oral verticils, each composed of about 1 6 tentacles. The tentacles of the row

40

MEDUS.E OF THE WORLD.

near the mouth are only about half as long as those of the lower circlet. In addition to these
there are about 25 long, tapering, hollow tentacles in a zone surrounding the base of the
polypite. During the breeding season, which occurs from January until May, great numbers
of medusae are developed upon the sides of the polypite immediately above the circlet of
basal tentacles. The budding medusae arise singly from the sides of the polypite and are not
produced in clusters upon peduncles as in Hybocodon christina, H. chilensis, and H. pulcher.
Longitudinal bands of orange pigment extend up the stem of the hydroid. The entoderm

FIG. 10. — Hydroid, young, and budding medusse of Hybocodon prolifcr, after L. Agassiz, in Con. Nat. Hist. U. S., vol. 4.

(A.) Mature hydranth. (B.) Oral eltremity of hydranth. (C.) Dissection showing intertentacular zone of medusa-buds.

Remaining figures are of medusa-buds and young medusae.

of the polypite is orange. L. Agassiz has shown that the entoderm of the stem is thrown
into longitudinal ridges which form partial septa projecting into the cavity of the stem. The
cavity, however, is continuous, and the septa do not fuse as in some other Hyboconidae.

This species is found upon the New England coast. According to Browne, 1896, it is
found upon the British, and also on the Irish coast, at Valencia, and off Iceland (Steen-

ANTHOMEDUS.E — HYBOCODON. 41

strup) and Norway (Sars). It is apparently widely distributed over the North Atlantic,
along the shores of continents and islands. It is rarely taken far from some coast. We
can not be certain that the American and European forms are identical until the hydroids of
both are discovered.

Hargitt, 1904, states that the egg-cleavage is closely similar to that of Pennaria. Also
Browne, 1895, and Hargitt, 1902, 1904, find that the ova begin their development within
the walls of the manubrium of the medusa and are set free as actinulae. Muiler, 1908, and
Hargitt, 1904, find that the developing embryos within the walls of the manubrium absorb
their fellow ova, as has been observed by Doflein in Tuhularia mesembryanthcmurn, and by
Allen in T. crocea. Developing actinulae and budding medusae are abundant at Woods Hole,
Massachusetts, during the spring months.

Hargitt, 1902, and Perkins, 1904, find that actinula larvae develop upon the manubrium
at the same time that medusa-buds are being set free from the tentacle-bulbs. When set free
the actinulae have 10 tentacles. The mouth and oral zone of tentacles develop only after the
actinula is set free; and appear at the pole which was adjacent to the parent medusa during
the attached period. Linko, 1905, also observed this simultaneous process of development
of medusa-buds and of actinula larvae in his " Amphicodon graviJum," which develops
actinulae with 1 1 tentacles. It is probably identical with H . prolifer. A single specimen was
found in Barents Sea, north Russia.

H. Miiller, 1908, finds that the ova are large and amoeboid. Only about 2 eggs survive
to maturity in the ovary; the others having been devoured by the successful eggs. The
ooplasma is a network, the exoplasma being narrow-meshed and the endoplasma wider.
There are numerous pseudo-cells in degenerate stages, sometimes dividing amitotically.

Hybocodon pendula Haeckel.
Plate i, fig. 2.

Corymorpha nutans (hydroid), STIMPSON, 1853, Marine Invert. Grand Manan, p. 9.

Corymorpha pendula (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 276, 343, plate 26, 6gs. 7-17. — VERRILL,

1873, Invert. Anim., Vineyard Sound, pp. 510, 736, plate 36, fig. 273.
Corymorpha pendula, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 85, fig. 81. — NUTTING, 1 901, Bull. U.S. Fish Commission,

vol. 19, pp. 337, 370, fig. 15. — MAY, 1903, American Naturalist, vol. 37, p. 579, n figs, (histology and embryology).
Corymorpha pendula (medusa), AGASSIZ, A., 1865, North Amer. Acal., p. 192, fig. 324.
Hybocodon pendula, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 34, plate 2, fig. 3.
Hybofodon pcndulus (medusa), HAECKEL, 1879, Syst. der Medusen, p. 34.
Monocautus pendulus, ALLMAN, 1871, Monograph Tubularian Hydroids, p. 397.

Adult medusa. — Bell pyriform and about 5 mm. in height. It is relatively higher than
the bell of H. prolifer, and the gelatinous substance at the apex is much thicker. There
are 5 rows of nematocysts upon the exumbrella, as in H. prolifer. The basal bulb of the
well-developed tentacle is much smaller than in H. prolifer, and no medusa-buds have
been observed to arise from it. The well-developed tentacle is 2 to 3 times the length of the
bell-height. Its surface is studded with large, swollen rings of nematocysts, which give it
a heavy appearance. The tentacles at the bases of the 2 intermediate radial-canals are quite
well developed, and this is not the case in H. prolifer. The velum is wide and thin. The
radial-canals are narrow and straight. The manubrium is longer than in H. prolifer, and
extends a short distance beyond the velar opening. There is a small peduncle. The lips
are thickly covered with nematocysts. Pink pigment-granules are found in the entoderm
of the tentacle-bulbs. The entoderm of the manubrium is pink and lilac, and contains also
some pink pigment-granules.

Hydroid. — The hydroid (Corymorpha[Hybocodori\ pendula)\s found in depths of IO to IOO
fathoms off the New England coast, with its base buried in the sand. It is 80 to 125 mm. in
height, and 6 mm. in diameter at the widest part. It always grows singly and is never branched.
The mid-region of the stem is very thick and is covered with a canaliculated coenosarc, but
the basal end narrows considerably, as does also the region near the free upper extremity,
which is long, slender, and pendulous. The stem is anchored by a number of root-like,
tubular, fleshy processes. The perisarc exists only as a thin delicate film. The head of the
polypite is large and highly contractile. There is a single verticil of long, hollow tentacles
at base of polypite. The mouth is situated at the extremity of a large flask-shaped proboscis,

42 MEDUSAE OF THE WORLD.

and is surrounded by a couple of rows of numerous irregularly arranged tentacles. These
oral tentacles are highly contractile, and are much smaller than those at the lower base of the
polypite. The medusae are borne upon branched stolon-like diverticula of the side walls of
the polypite, immediately above the zone of basal tentacles. Fully-developed medusae have
not been seen to be set free from the hydroid, but the similarity of the most advanced medusa-
buds observed to the free medusa found in the ocean leaves but little doubt concerning this
point. May, 1903, has studied the histology and embryology, and concludes that the medusa-
buds may at times become free, but usually mature while still attached to the hydranth.
This species has been found from Vineyard Sound to the mouth of the St. Lawrence River.
The medusa appears upon the southern New England coast in April and May, but is not seen
during the summer months, although the hydroid is abundant at this time.

It is possible that Euphysa tentaculata Linko, 1905, is identical with H. pendula (see
Zool. Anzeiger, Bd. 28, p. 214). Linko's medusa is from Barents Sea, north of Russia. I
have referred to this medusa in the description of Steenstrupia bigelowt.

Hybocodon forbesii Mayer.
Plate I, fig. 8; plate 2, fig. 3.

Hybocodon jorbesii, MAVF.R, 1894, Bull. Mus. Comp. Zool. at Harvard College, vol. 25, No. II, p. 236, plate I, fig. l; 1904,
Memoirs Nat. Sci. Museum Brooklyn Institute, vol. i, No. i, p. 8, plate 2, fig. 13.

Medusa. — Bell asymmetrical, about 2.5 mm. in height and ellipsoidal in shape, being
slightly higher than broad. The gelatinous substance is of uniform thinness. There is a
single well-developed tentacle situated at the base of the longest radial-canal. A short, conical
tentacle is found at the base of the shortest radial-canal, and two smaller tentacle-bulbs are
situated one at the base of each of the intermediate canals. The well-developed tentacle is
about as long as the bell-diameter. Its base is small, and hardly greater in diameter than
the shaft of the tentacle. Its free extremity is fusiform, and covered with prominent nemato-
cyst-cells. No medusa-buds have ever been observed. The velum is narrow. The 4 radial-
canals are straight and slender and the circular canal is narrow. There are no rows or clusters
of nematocysts upon the exumbrella. Manubrium is spindle-shaped and swollen, and the
mouth is a simple, round opening situated at the extremity of a narrow tubular neck, which
extends beyond the velar opening. The entoderm of the terminal swelling of the large ten-
tacle is yellow streaked with red. Entoderm of manubrium yellow with red flecks.

This species is found in Nassau Harbor, New Providence Island, Bahamas, and at
Tortugas, Florida, in March to May. It is an abundant surface form. I have captured
many hundreds of specimens, but have never found them producing either medusa-buds or
actinula larvae. It is distinguished by its decided yellow and orange color, and the absence
of meridional lines of nettle-cells over the exumbrella.

"Hybocodon unicus."

Amphicodon unicus, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

(?) Hybocodon chilensis (hydroid), HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545, fig. W.

Bell 3 mm. high, 2 mm. wide, bell-shaped. I solitary tentacle between 2 rudimentary
basal bulbs; 3 perradially situated bulbs without tentacles. Medusa-buds ( ?) Manubrium
cylindrical, nearly as long as the umbrella cavity. Color ( ?) Asymmetry of bell ( ?) One
specimen was found by Vallentin, and briefly mentioned without figures by Browne, from
Stanley Harbor, Falkland Islands. Hybocodon chilensis Hartlaub, from the coast of Chile,
may prove to be the hydroid of this medusa ( ?) It will be impossible to identify the medusa
from the brief mention of it given by Browne, unless, indeed, it be rediscovered in Stanley
Harbor.

Hybocodon chilensis Hartlaub.

Hybocodon chilensis, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545, fig. W.

(?) Amphicodon unicus (medusa), BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

(?) Steenstrupia occidentalis (medusa), FEWKES, 1889, Bulletin Essex Institute, Salem, vol. 21, No. 7, p. 107, plate 3, fig. I.

Hybocodon occidenlalis, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545.

As Hartlaub states, this hydroid may be the stock of Amphicodon (Hybocodon) unicus
Browne, from the Falkland Islands.

ANTHOMEDUS.E — HYBOCODON. 43

The hydrocaulus is very thick and massive, about 50 mm. long and unbranched save for
the presence of" its roots and stolons. It is covered with a stiff layer of chitin, which is not
expanded at the base of the polypite. Polypite large, with 17 to 20 proximal tentacles, each
about 6 mm. long. There are also about 27 oral tentacles arranged in several rows. Above
the bases of the proximal tentacles there are 8 long, thick, medusa-bearing stolons, which
are thickly covered with numerous clusters of medusa-buds. Each medusa-bud has a single
very large tentacle. The stem of the hydroid is rusty yellow and the polypite light rose-color.
Found at Calbuco, Chile, South America.

This Hybocodon is closely related to, or possibly identical with, the form from Norway
described by Bonnevie under the name Hybocodon prolifer, but it differs from H. prolifer
Agassiz in having large, specialized, medusa-bearing stolons. Hartlaub proposes to call this
Norwegian hydroid Hybocodon christincc.

Fewkes, 1889 (Bull. Essex Inst., and also Amer. Naturalist, vol. 32, p. 597), gives a
brief description of a medusa from the coast of California which he calls by two names, Steen-
strupia occidentahs and S. californica; and which may be derived from Hybocodon chilensis.
This medusa is described as follows: Size (?) Bell ovoid without an apical prominence.
4 (?) 5 ( ?) rows of meridional lasso-cells extend upward from the 4 tentacle-bulbs, over the
exumbrella toward the bell-apex. I long tentacle and 3 rudimentary tentacle-bulbs at the
bases of the 4 radial-canals. The long tentacle is ringed at regular intervals and has a
large pigmented basal bulb from which there arise numerous medusa-buds. Color ( ?)
Velum well developed. 4 straight, narrow radial-canals. Manubrium shorter than the
depth of the bell-cavity. No axial canal. Coast of California, United States.

Hartlaub proposes to call this medusa Hybocodon occidentalis. I find nothing in Fewkes's
description to distinguish it from H. prolifer L. Agassiz, but apparently there is less difference
between the medusas of the various forms of Hybocodon than between their hydroids.

Hybocodon christinae Hartlaub.

Tubular ia prolifer^ Hybocodon prolifer, BONNEVIE, 1899, The Norwegian North Atlantic Eipedition, 1876-79,70!. z6, Hydroida,

p. z8, plate I, fig. 6.
Hybocodon chrislin,t, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 546; 1907, Nordisches Plankton, Nr. iz, p. loz, fig. 98.

The medusa attributed by Bonnevie to this hydroid has a single well-developed tentacle
with a basal cluster of large medusa-buds, each bud bearing a single tentacle. The medusa-
buds resemble H. prolifer, but the hydroid is distinguished by bearing its medusae upon 8
branched peduncles.

Hydrocaulus unramified, tubular, springing from a ramified hydrorhiza; occurrence
solitary; longitudinal striping, no collar; height about 50 mm. The hydranth has 14 proxi-
mal tentacles and 2 distinct circles of (oral) distal tentacles. The oral tentacles are shorter
and more numerous than the proximal. There are 8 blastostyles in a circle about midway
between the oral and basal tentacles, and these bear numerous medusae upon short pedicels.
The medusa-buds have 4 very wide radial-canals, and I highly developed tentacle which
exhibits at its swollen base the bud-rudiments of 4 new medusae even before the first has
become detached. This species is distinguished by its well-developed, branched, medusa-
bearing stolons. Found off Bodo, Norway. It is closely related to H. chilensis Hartlaub, of
the northern coast of Chile.

The medusa-buds in H. christinie appear to be confined to the under side and the sides
of the base of the well-developed tentacle.

Hartlaub, 1907, finds that the medusa becomes 4 mm. high and 3 mm. wide, with thin
bell-walls and an evenly rounded apex. The 4 radial-canals and ring-canal are band-like,
and wider than in other species of Hybocodon.

Hybocodon pulcher Hartlaub.

Auliscus pulcher, SAMUNDSSON, 1899, Vid. Meddcl. Nat. For. Kjobenhavn, Ser. 6, Aarg. i, p. 4Z5, taf. 4, 7 figs.

Hybocodon pulcher, HARTLAUB, 1905, Zool. Jahrbuchern, Suppl. 6, p. 545 ; 1907, Nordisches Plankton, Nr. iz, p. 96, figs. 9Z, 93.

Hydroid 40 to 50 mm. high with an oral circlet of 30 short tentacles, and another circlet
of 24 to 30 long tentacles around widest part of body of the hydranth. The only distinctive
characters of this species are the well-developed medusa-bearing stolons, the symmetrical

44

MEDUSAE OF THE WORLD.

bell, and 2 large principal tentacles of the budding medusa. In H. prolifer, on the other hand,
the medusae are usually set free with but one well-developed tentacle. The bell of H. pulcher
may become asymmetrical in later life ( ?) The hydrorhiza is thin and branching. There is a
flexible collar-like pensarc at the base of the hydranth. Medusas are developed upon stolons
from the hydranth above the circlet of long tentacles. When set free the medusa is 1.5 to 2 mm.
high, of symmetrical form. 5 longitudinal lines of nematocysts extend up the sides of the
bell to the apex. 3 small tentacle-bulbs. 2 equally well-developed tentacles arise side by
side, from the base of one of the radial-canals. Entoderm of manubrium and tentacles red.
Found off Iceland. I believe this form is probably identical with H. prolifer.

Genus MICROCAMPANA Fewkes, 1889.

Microcampana, FEWKES, 1889, Amer. Naturalist, vol. 32, p. 595; Bull. Essex Inst., Salem, vol. 21, No. 7, p. in.

The type-species is Microcampana conica Fewkes, from Santa Cruz Island, off the coast
of California.

GENERIC CHARACTERS.

Anthomedusae with 6 radial-canals, and 6 radially placed marginal tentacles. One of
these tentacles is well developed, and the other 5 are rudimentary.

It is possible that the vaguely described Rhabdoon singulare, of Keferstein und Ehlers,
(1861, Zoolog. Beitrage, p. 86, taf. 13, figs. 6, 7), from Messina, Mediterranean, is a form
of Microcampana, but there are apparently 12 longitudinal lines of nettling cells over the
exumbrella, and it is uncertain whether there are 4 or 6 radial-canals. The bell is 1.5 mm.
high and oval with uniformly thin walls. It is possibly an abnormal medusa of P elella.

Microcampana conica Fewkes.

Microcampana conica, FEWKES, 1889, Bull. Essex Inst., Salem, vol. 21, No. 7, p. in, plate 4,
fig. 8; American Naturalist, vol. 32, p. 595, fig.

Size ( ?) Bell conical with a well-developed, elongate, conical
apex; slightly asymmetrical. Exumbrella smooth, without meridional
rows of nematocysts. 6 marginal tentacles, 60° apart. 5 of these
are rudimentary, but the sixth is club-shaped, and about half as long
as the bell-height. 6 straight, narrow radial-canals and a ring-canal.
Manubrium conical to spindle-shaped, about as long as the depth of
the bell-cavity. There is a long, slender, straight, axial canal above
the stomach. Bell pink, tentacle-bulbs bright-red, manubrium yellow.
Found off Santa Cruz Island, California; under the cliffs of Punta
Diablo.

Genus DICODONIUM Haeckel, 1879, sens. ampl.

Dieodonium+Dinemas HAECKEL, 1879, Syst. der Medusen, pp. 27, 28.

ATorc Dinema, VAN BEN ED EN, 1867, Mem. Acad.Roy.des Sci.Belgique, 10111.36, art. 2, pp. 127, 130.

Dicodonium, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 443.

Sarsiella, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 66.

The type-species of this genus is DicoJonium cornutum Haeckel, 1879, of the Red Sea.
Dinema Van Beneden is a medusa which arises by budding from a Perigotiirnus-\\ke hydroid
and therefore belongs to the Tiannae. We use the term Dicodonium in the sense proposed
by Vanhoffen, 1891.

GENERIC CHARACTERS.

Codonidae with 2 well-developed and 2 rudimentary tentacles. No meridional lines of
nettle-cells upon the exumbrella. An apical projection to the bell and an axial canal pro-
jecting upward from the stomach, may or may not be present.

Some of the so-called "species" of Dicodonium are probably only abnormal specimens
of Sarsia with 2 tentacles instead of the normal 4.

FIG. II. — Microcampana
after Fewkes in Amer-
ican Naturalist.

PLATE 3.

Fig. I. Dicodonium Jefferson:. Tortugas, Florida, June 15, 1897.

Fig. 2. Sarsia rnirabilis, young medusa. Nahant, Massachusetts, March

26, 1897.
Fig. 3. Hybocodon prolijer. t\ primary; t", secondary tentacle beginning to

develop; m, young medusa-buds. Agassiz Laboratory, Newport,

Rhode Island, September 17, 1895.
Fig. 4. Sarsia mirabilis, mature male. Nahant, Massachusetts, May 7,

1897.
Fig. 5. Hydroid of Sarsia mirabilis with ripe male medusa-bud attached to

the hydranth. Swallows Cave, Nahant, Massachusetts, May 8,

1897.

Drawn from life, by the author.

PLATE 3

ANTHOMKDUS.E — DICODONIUM.

45

Tabular Description of the Medusa of DicoJonium.

D. cornutum
Haeckel.

D. dissonema
Haeckel.

D. ocellatum
= Dinema
ocellatum
Haeckel.

D. floridana
Mayer.

D. jeffersoni
Mayer.

D.adriaticum
Graeffe.

D. dinema =
Sarsiella
dinema
Hartlaub.

Shape and size

With bulging

Bell-shaped to

Bell-shaped

Cylindrical with

Dome-like

Bell-shaped

Half-egg-

of bell in

sides, and

egg-shaped

to hemi-

dome-like

0.75 high,

with a small,

shaped. 3

mm.

pointed

with blunt

spherical. 5

apex. 4

0.5 wide.

conical, api-

high, 2 wide.

apex. 4

conical

high, 4 to 5

high, 3 wide.

cal projec-

high, 4 wide.

a prx.

wide. Ex-

tion. 4 high,

umbrella

3.5 wide.

besprinkled

with tufts of

nettle-cells.

Length of 2

Curled up-

6 r ± lone,.

2 r lun^. v. irli

2 r long with a

r long. Basal

8 r long. A

2 r + long.

long tenta-

ward. 3 r

Verv large

fairly large

knob-like

bulbs bear

large car-

Basal bulb-

cles m

long. A

basal bulbs

tentacle-

swelling near

ocelli.

mine ocellus

small; with

terms of

row of bns-

and with an

bulbs, each

tip. Extreme

on each ten-

.'i .'Hi.

bell-radius

thngclusters

abaxial ocel-

with an ab-

tip ends in

tacle-bulb.

«.

of nettle-

lus.

axial ocellus.

thin lash. No

cells on their

ocelli.

abaxial

sides. Tips

club-shaped.

Basal bulbs

small.

Condition of

Not present.

>

Mere bulbs.

Small, taper-

Mere basal

Men- basal

Absent.

2 rudimen-

ing, rudimen-

bulbs with

bulbs with

tary tenta-

tary.

abaxial

small car-

cle-bulbs.

ocelli.

mini- "1 rill.

Axial canal

Well devel-

Present.

Not present.

Not present.

A small axial

Not present.

above stom-

oped .

canal is

ach.

presrnt.

Shape of man-

Spindle-shap-

Spin. He-shap-

Club-shaped.

Flask-shaped,

Cylindrical,

Short, thick,

Spindle-

ubrium,

ed. 1.5 r

ed. 2 to 3

5 to 7 r

widest above,

with a nar-

and four-

shaped,

and length

long. Mouth

times as

long.

but near mid-

row tubular

sided.

6r long.

in terms of

a simple

wide in mid-

dle.

mouth-end.

bell-radius

round open-

dle as at

«•

ing sur-

either end.

rounded bv

nettle-cells.

Gonail.

Ring-like, swol-

Large swollen

Gonad en-

Encircling

Large gonad

In stomach

Encircles en-

len, encir-

gonad en-

circles stom-

stomach.

encircles

wall.

tire stom-

cling middle

circles stom-

ach, thickest

stomach

ach.

third of

ach.

near mouth.

from base to

manubrium

near mouth.

Color.

Stomach, ten-

)

Bell-margin

Entoderm of

Entoderm of

p

Entoderm

tacles, and

rose-re, 1.

stomach and

stomach

brownish-

nettle-ring

Nematocyst

tentacles yel-

creamy

yellow.

of margin

clusters of

low. Knoh-

pink; of

dark purple-

exumbrella

like ends of

tentacle-

red.

black.

tentacles

bulbs deli-

Ocelli of ten-

flecked with

cate green

tacle-bulbs

red to orange.

or pink.

black.

Ocelli red.

Where found.

Red Sea, Gulf

Coast of Aus-

Trieste, Adria-

Tortugas, Flor-

Tortugas, Fla.

TruMr, Adria-

Normandy,

of Suez.

tralia.

tic Sea.

ida. On sur-

M.iv to

tic Sea.

coast of

face in June

June, com-

France.

to July, rare.

mon.

Mediterran-

ean ?

Remarks.

Development

Development

Development

Development

Development

1 ' • 'pment

unknown.

unknown.

unknown.

unkri"u n.

unknown.

unknown.

Is this an

Occasionally

•

This species

abnormal

rudimentary

is distin-

Sarsia with

tentacles de-

guished by

only 2 ten-

velop so as to

stiff sensory

tacies ?

be almost as

hairs which

long as the

border ocelli

larger pair.

46 MEDUSAE OF THE WORLD.

Dicodonium cornutum Haeckel.

Dicodonium cornutum, HAECKEL, 1879, Syst. der Medusen, p. 27, taf. I, fig. 6.

Haeckel found this species at Tur, near Sinai, in the Red Sea. See tabular description of
medusae of Dicodonium. It has no ectodermal ocelli upon the bulbs of the 2 large tentacles.

Haeckel presents a beautiful figure of the medusa, drawn
from life.

Dicodonium dissonema Haeckel.
Dicodonium dissonema, HAECKEL, 1879, Syst. der Medusen, p. 27.

Haeckel describes this from a preserved specimen from
the coast of Australia. See tabular description of the
medusae of Dicodonium.

"Dicodonium ocellatum."

FIG. 12. — Dicodonium cornutum, after Sarsia ocellata, BUSCH, 1851, Beobach. wirbellos. Seeth., p. 16, taf. 2, figs. 1-3.
Haeckel, 1879. Dinema ocellatum, HAECKEL, 1879, Syst. der Medusen, p. 29.

This medusa is described by Busch from Trieste, Adriatic, and it is probably an abnormal
Sarsia with only 2 of its 4 marginal tentacles developed. See tabular description of the medusae
of Dicodonium.

Dicodonium floridana Mayer.
Plate 2, fig. 5.

Bell about 4 mm. high and 3 mm. wide, with thin, uniform, vertical walls and a slight
apical projection. There are 2 equally-developed, diametrically opposed tentacles, each
about three-fourths as long as the bell-height. Near the outer end of each of these tentacles
there is a large, knob-like, swollen region, which terminates in a thin, tapering, nematocyst-
bearing lash. The knob-shaped swelling is hollow and its cavity is connected with the general
gastrovascular system of the medusa by means of a slender tube which extends through
the entodermal core of the tentacle. The basal bulbs of the tentacles are not large, and there
are no ocelli.

In addition to the 2 long tentacles, there are 2 small, tapering, rudimentary tentacle-
bulbs 90° from the large tentacles. The velum is well developed. There are 4 straight,
narrow radial-canals and a simple, narrow circular canal. The manubrium is flask-shaped,
being narrower at its base than at its middle point. The mouth projects slightly beyond the
velar opening, and is a simple, round opening at the extremity of a long, tapering neck. The
gonads are within the wall of the manubrium. The entoderm of the stomach is yellow, and
that of the distal bulbs of the tentacles yellow flecked with orange. The entoderm of the basal
bulbs of the tentacles is also tinged with yellow.

This medusa is occasionally found at Tortugas, Florida, in June. Occasionally a speci-
men is taken in which the normally rudimentary tentacles have developed so as to be nearly
as long as the pair of large tentacles, thus illustrating the imperfect line of separation between
Dicodonium and Sarsia.

Dicodonium jeffersoni Mayer.

Plate 2, fig. 4; plate 3, fig. I.
Dinema jeflersoni, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 30, plate 37, fig. 126.

The bell is dome-shaped, higher than a hemisphere, and about 0.75 mm. high. The
exumbrella surface is sparsely sprinkled with nematocysts. There are 2 short marginal
tentacles, and 2 rudimentary tentacle-bulbs. The tentacles are radially situated, and are
covered with numerous small wart-like clusters of nematocysts. There are 4 ectodermal
ocelli, I upon the outer side of each of the 4 tentacle-bulbs. The velum is well developed.
There are 4 straight, narrow radial-canals and a narrow circular vessel. The manubrium
is about as long as the depth of the bell-cavity. It is simple, cylindrical, and tube-like, and
the mouth is a round opening at the extremity of a short, cylindrical neck. A simple canal

ANTHOMEDUS^E — DICODONIUM, SARSIA. 47

projects upward from the stomach into the gelatinous substance of the apex of the bell.
This is probably only the remnant ot the connection between the medusa and its hydroid
stock. The gonad is ring-like, and encircles the stomach, leaving the short proboscis free.

The entoderm of the tentacles and tentacle-bulbs is of a delicate green or pink. The
entoderm of the stomach is creamy pink. The ocelli are bright-red, and all other parts are
colorless.

This medusa is quite common at the Tortugas, Florida, in May and early June. Although
small, it appears to be mature, for sperm is often given off from the gonad of the males.

Dicodonium adriaticum Graeffe.
Dicodonium adriaticnm, GRAEFFE, 1884, Arbeit. Zool. Inst. \Vien, Bd. 5, p. 551.

Bell 4 mm. high, 3.5 mm. wide, bell-shaped, with a small, conical, apical projection.
2 long, radially placed tentacles, more than 12 mm. long, with large, thick, basal bulbs, each
with a large carmine ocellus upon the abaxial side of the bulb. 2 tentacle-bulbs at the bases
of the 2 radial-canals 90° away from the large tentacles. These basal bulbs have small red
ocelli. Each of these 4 ocelli are bordered by a ring of stiff sensory hairs. There are 4 small,
interradial tentacle-bulbs without ocelli. Thus there are 2 long tentacles, and 6 rudimentary
tentacle-bulbs. 4 radial-canals. Stomach short, thick, and 4-sided, with the gonads in the
stomach-wall. Mouth simple, with 4 lips. Found at Trieste. Adriatic Sea, in October.

Dicodonium dinema.
Sarsiclla dintma, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 67, fig. 63.

Bell oval, half-egg-shaped, 3 mm. high, 2 mm. wide. Exumbrella thickly besprinkled
with nematocysts. Only 2 tentacles, 1 80° apart. These are longer than the bell-diameter.
They have small basal bulbs, each with a reddish-brown ocellus. No trace of tentacle-bulb
or tentacles 90° apart from the well-developed tentacles. Manubrium about 2 times as long
as the bell-height. Spindle-shaped and encircled throughout by the gonad. Manubrium,
tentacles, and 4 radial-canals brownish-yellow. Found off the coast of Norway and in the
Mediterranean ( ?) Is this an abnormal young Sarsia with only 2 tentacles ? Hydroid
unknown. Medusa rare.

Genus SARSIA Lesson, 1843.

Sarsia, LESSON, 1843, Hist. Zooph. Acal., p. 333. — AGASSIZ, L., 1849, Mem. Amer. Acad., New Scries, vol. 4, p. 224; 1862, Cont.
Nat. Hist. U. S., vol. 4, p. 211. — WAGNER, 1885, Wirbellosen des Weissen Meeres, p. 76. — VANHOFFEN, 1891, Zool.
Anzeiger, Bd. 14, p. 442. — HARTLAUB, 1897, Hydromedusen Helgolands, p. 454; 1907, Nordisches Plankton, Nr. 12,
p. 7. — VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 582. — CHUN, 1895, Bibliothcca Zoologica,
Bd. i, Heft. 19, p. 4. — GOETTE, 1904, Zool. Anzeiger, Jahrg. 27, p. 473.

Coryne + Syndictyon, AGASSIZ, A., 1865, North Amer. Acal., p. 175.

Coryne (hydroid), CALKINS, 1899, Proc. Boston Soc. Nat. Hist., vol. 28, p. 336.

Codonium + Sarsia + Svndictvon, HAECKEL, 1879, Syst. der Medusen, pp. 13, 16, 20.

Syncoryne, WEISMANN, 1883, Entsteh. Sexualzellen Hydromedusen, pp. 56, 216.

Syncoryne (hydroid), HARTLAUB, 1905, Zool. Jahrbuchern, Suppl. 6, p. 524. — NUTTING, 1901, Proc. Washington Acad. Sci.,
vol. 3, p. 165.

Syncoryne (medusa), HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, pp. 29, 30.

Syncoryne, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 119 (citation of all references to 1850).

SynJiclon, AGASSIZ, A., 1862, in L.Agassiz's Cont. Nat. Hist. U. S., vol. 4, p. 340. — MAYER, 1904, Mem. Nat. Sci. Museum Brook-
lyn Institute, vol. i, p. 7.

Stenyo, DUJARDIN, 1845, Annales Sci. Nat., ser. 3, tome 4, p. 257.

The type-species of this genus is Sarsia tubulosa of the northern coasts of Europe. This
medusa was first described by Lesson, 1843. The hydroid form was first described by
Gartner, 1774, in Pallas's Eleunch. Zooph., under the name of Coryne. Ehrenberg, Sars,
and Allman introduced the name Syncoryne. Staurtdia producta also gives rise to a medusa
which can not be distinguished from Sarsia.

GENERIC CHARACTERS.

Codonidae with 4 long, simple, equally developed tentacles, I at the foot of each radial-
canal. The manubrium is tubular and surrounded by a ring-like gonad. There is an
ectodermal ocellus upon the outer side of each tentacle-bulb. There are no meridional
nematocyst-tracts upon the exumbrella.

48 MEDUSA OF THE WORLD.

An apical projection of the bell may or may not be present; and there may or may not be
an axial canal extending upward from the stomach into this projection. The hydroid is
Syncoryne or Staundia.

We use the name Sarsia in the sense defined by Vanhoffen, 1891 (Zool. Anzeiger, p. 442).

In 1862, 1865, A. Agassiz described, under the generic name Syndictyon, a Sarsia having
reticulate nematocyst-cells upon its exumbrella, and clusters of such cells upon its tentacles.
These are, however, only characters of immaturity and largely disappear in the full-grown
medusa, which is a true Sarsia in all respects. In 1879, Haeckel formed the genus Codonium
to include medusae resembling Sarsia but distinguished by the possession of an apex upon
the bell into which a blindly-ending axial canal extends from the stomach. A bell-apex
and axial canal are characters which are acquired during growth in varying degrees by
almost all species of Sarsia, and are therefore not of generic value.

Two European and one American species of Sarsia produce medusae by asexual budding
from the tentacle-bulbs or from the walls of the manubrium. Chun, 1895 (Bibliotheca Zoo-
logica, Heft 19, fig. 2), showed that both ectoderm and entoderm of the manubrium take
part in the formation of these proliferating medusae; the entoderm of the manubrium forming
the entoderm of the daughter medusa, and the same being true of the ectoderm. Sarsia
prolifera Forbes, described and beautifully figured by Haeckel, 1879, under the name Codonium
codonophorum, produces medusa-buds upon its tentacle-bulbs.

The majority of Sarsia medusae are probably produced asexually by hydroids of the
genus Syncoryne, but at least one medusa identical with Sarsia is derived from the hydroid
called Staundia Dujardin, 1843. Such medusae may conveniently be placed in a subgenus
Staundwsarsia. A medusa which appears to be closely related to Sarsia is produced by the
remarkable parasitic hydroid Hydrichth vs.

The generic name Syncoryne was restricted by Allman, 1871-1872, to designate the
hydroid which produces the medusa Sarsia. The name Syncoryna was first proposed by
Ehrenberg, who applied it to hydroids now known as Clava, Coryne, etc., and in this old
sense it does not apply exclusively to the hydroid of Sarsia. By general consent, Allman's
name has been accepted in this restricted sense, Calkins, 1899, being almost alone in main-
taining that the generic name of the hydroid should be Coryne.

The commonly accepted arrangement is to retain the old name Coryne to include hydroids
in which the reproductive elements are produced in fixed sporosacs growing upon the
hydranth, while Syncoryne applies to like hydroids which, however, produce free medusae.

Weismann, 1883, found that the germ-cells of both sexes of Syncoryne sarsii originate
in the ectoderm of the budding medusa, and do not wander from their place of origin, but
become mature in the free medusa. Goette, 1904, finds, however, that in Sarsia the sperm
originates and remains in the peripheral ectoderm of the manubrium of the medusa, but the
pgg-cells, contrary to Weismann's contention, originate in the entoderm of the medusa-bud
while it is still attached to the hydroid, but afterwards they migrate into the ectoderm of
the manubrium, where they mature.

Many of the species of Sarsia display considerable individual variability, the colors of
the manubrium and tentacle-bulbs ranging from green or yellow to red. An apical pro-
jection and an axial vessel above the stomach may or may not be developed, and the length
of the manubrium at maturity is subject to much variability. Moreover, the hydroids may
form densely or sparingly branched colonies in accordance with environmental conditions,
and as is well known in S. mirabihs, free medusae are produced in early spring, whereas late
in the season the medusae mature while still attached to the hydroids. Much contusion has
been introduced into the synonymy of the genus, and different stages of the same medusa
have occasionally received different specific names.

Sarsia " nodosa" Busch, 1851, appears to be the young of some European Sarsia. The
exumbrella of very young medusae of Sarsia are usually besprinkled with nettling cells and
their tentacles bear prominent nematocysts, and Busch's medusa displays both of these
characters. (See Hartlaub, 1907, Nordisches Plankton, Nr. 12, p. 68, fig. 64.)

ANTIKIMKDrs.K — SAKSIA.

49

Tubular Description of the Medusa of Sarsia.

S.tubulosa Lesson,

i843*.

S. mirabilis L. Agassiz
•»49-t

S. eximia = Syncor-
yne eximia
Hincks, 1868.

S.radiata von
Lrndenfeld.

S. conica= Codon-
ium conicum

Haeckd.

Shape and size of

Egg-shaped. Half-

As in S. tubulosa, 7 to

Oval with quite

Semi-ovate. 3

Barrel-shaped with

bell in mm.

ellipsoidal. loto

10 high, 4 to 8 wide.

thick walls. 3

high, 2. 5 wide. long p<

12 high, 6 to 8

high. 2 wide.

apex. 12 high, 4

wide. With or

wide.

without apical

projection.

Character of 4 ten-

Basal bulbs small;

As in S. tubulosa.

Basal bulbs large,

As in S. eximia

7 r long, with small

tacles. Length

each with abaxial

with abaxial ocel-

basal bulbs.

in terms of boll-

ocellus. Tentacles

lus on each. Ten-

radius (r).

about 4 to 6 r

tacles about 4 r

long, and without

long-

prominent clusters

of nettle-cells.

Shape and length

Narrow, cylindrical

Narrow tubular base.

Cylindrical; wide

As in S. eximia,

Spindle-shaped,

of manubnum

at its base. Mid-

Mid-region swollen and

above, with nar-

i .5 r long or

only 1.5 to 2 r

in terms of bell-

dle part cylindri-

cylindrical. Line of

row, short, tubular

less.

long.

radius (r).

cal, swollen by

demarcation between

neck above mouth

gonad. Stomach

narrow, cylindrical,

Length about 2 r

small, spindle-

basal, and wide middle

or less.

shaped. Mouth at

part very sharp.

end of short coni-

Stomach swollen and

cal neck, as in

spindle-shaped and

S. mirabilis.

near outer end of tin-

Length 4 r.

manubrium. Beyond

stomach there is a

short, tapering, narrow

neck. Mouth is a

round opening. Total

length 3 to 6 r. Average

4 r.

Gonads.

Ring-like and de-

Thick ring encircling

Encircles manu-

As in S. eximia.

Encircles stomach,

veloped over

manubrium. Begin-

brium from base

leaving both ends

nearly the whole

ning abruptly at a short

to mouth end,

free.

length of the

distance from base,

leaving only short

manubnum, leav-

and extending to upper

neck above the

ing a short dis-

part of stomach, leav-

mouth free. No

tance at both ends

ing both ends of man-

medusa-buds.

free. No medusa-

ubnum free.

buds.

Color.

Entoderm of stom-

Entoderm of stomach

Entoderm of stom-

Entoderm of

?

ach green, bluish

usually green, some-

ach yellowibh-

manubrium

lilac, or red-

times green and red,

red to reddish-

and tentacle-

Tentacle-bulbs

or red. Tentacle-bulbs

brown or green.

bulbs deep

green, blue, or red.

green or red. Ocelli

Ocelli black in

brown.

Ocelli black.

black.

young and ml m

mature medusa.

TentacJe-bulbs

red, or brownish-

red.

\Vhere found.

North Atlantic

From coast of New Jrr-

Coast of England,

New South

Indian (Kc.m.

coasts of Europe

scv northward to Arctic

S( ntland, Helgo-

W.ilrs, Aus-

to Arctic Ocean.

Ocean. Coasts of Nor-

land, Nnnvav,

tralia.

wav. northern Russia,

Juneau, Alaska,

Greenland, Alaska to

North Pacific

northern Chile along

(Nutting).

Pacific coast to South

America.

Hydroid.

Syncoryne sarsia

Svnotrvne "miraluli "

Svncoryne eximia,

Sviu i>rvnr r.uli-

1'iikrmwn.

Loven. Hartlaub,

L. Agassi/,. S. reticu-

Allman.

aba.

1905, describes

lata is a closely related

(this hvdroid ?)

variety (see text).

from Terra del

Fuego, Antarctic.

*Description based on accounts by European writers.

j-Description based on original observations of medusa: found off southern coast of New England, United States.

50

MEDUS/E OF THE WORLD.

A number of Syncoryne hydroids have been described which probably produce Sarsia
medusae, but are not known so to do. Among these may be mentioned Syncoryne crassa
Pictet, 1893 (Revue Suisse Zool., tome I, p. 8), a small hydroid only 2 mm. high and rela-
tively thicker and shorter than S. pulchella of Allman, 1871 (Monog. Tubularian Hydroids,
p. 279, plate 6, fig. 3). Pictet's hydroid comes from Amboina, Malay Archipelago. It has
30 to 40 short, knobbed tentacles, and the medusa-buds arise singly between the tentacles.
The hydrorhiza is net-like, and the hydroid is pale orange in color.

Hartlaub, 1907, has made a masterly study of the genus Sarsia, and shows that the
species fall conveniently into two groups: the Eximia group, with short manubrium occupied
entirely by the gonad, which extends from its base to near the mouth, and the Tubulosa group,

Tabular Description of the Medusa of Sarsia. — Continued.

S. prolifera Forbes
= S. codono-
phora Haeckel.
(See text.)

S. gemmifera Forb.
= S. siphono-
phora Haeckel.

S.hargitti = S.pro-
ducta Hargitt.

S. flammea Hart-
laub, 1907.

Sarsia (Stauridio-
sarsia)producta =
Stauridiaproducta
Wright, Hartlaub.

Shape and size of

Bell-shaped, with

Bell ellipsoidal.

Pyriform. 1.5 high,

Oval. 12 high, 7

Three-fourths-egg-

bell in mm.

or without apical

8 high, 6 wide.

i wide.

wide.

shaped. 10 high,

projection. 2.5

7 wide.

to 8 high, 3 to 8

wide.

Character of 4 ten-

Basal bulbs very

Well-developed

Well-developed

Tentacle tips knob-

Large basal bulbs

tacles. Length

large and wide,

basal bulbs with

basal bulbs with

like. Shafts cover-

with abaxial ocelli.

in terms of bell-

with black or red

abaxial ocelli.

abaxial ocelli.

ed with broken

Tentacles about 3

radius (r).

ocelli. Tentacles

Tentacles4r long.

Tentacles ir long.

(partial) rings.

to 4 r long.

hollow, tapering.

No ocelli.

2 to 6 r long;

with clusters of

medusa-buds

upon their bases.

Shape and length

Spindle-shaped with

Very long, tubular,

Spindle-shaped at

Conical — spindle-

Cylindrical, about

of manubrium

narrow neck and

with spindle-

both ends, narrow-

shaped, only two-

3 r long.

in terms of bell-

simple, round

shaped stomach

er and tubular in

thirds as long as

radius (r).

mouth-opening.

near outer end.

middle. 2 to 3 r

the depth of the

About i .5 r long.

Mouth at extrem-

long. A circlet of

bell-cavity.

ity of narrow

medusiform

neck. Manubrium

gonads( = medusa-

8 r long.

buds ?) near prox-

imal end of man-

ubrium.

Gonads.

Encircling stomach.

Gonad ( ?)

Gonad ( ?)

Ring-like, encirc-

Gonad at base of

Medusae pro-

Medusae produc-

Medusiform buds

ling the manu-

stomach. No me-

duced asexually

ed asexually by

on stomach in cir-

brium from its

dusa-buds.

upon the tentacle-

budding from

clet near base.

base to near

bulbs.

sides of manu-

mouth.

brium. There may

be 10 to 22 or

more of these

budding medusae

upon manubrium

at one and same

time. Law of bud-

ding discussed in

text.

Color.

Entoderm of tenta-

Entoderm of stom-

Basal part of manu-

Entoderm fiery red

Stomach brownish,

cles and stomach

ach, tentacles, and

brium orange.

or orange.

gonads yellowish-

yellow to sage-

radial-canals or-

Distal end blue-

white, tentacle-

green. Mouth or-

ange-yellow to or-

green. Tentacle-

bulbs red. Ocelli

ange. Ocelli

ange-red. Ocelli

bulbs orange,

black.

brown. Tentacle-

black.

edged with green.

bulbs contain red

Ocelli black.

entodermal pig-

ment.

Where found.

English Channel to

Atlantic coasts of

No Man's Land,

Arctic Ocean.

Coast of Great

Mediterranean.

Europe, from

near Woods Hole,

Britain to Helgo-

Norway south-

Massachusetts,

land.

ward to Canary

United States. A

Islands.

single specimen

was found.

Hydroid.

Unknown.

Unknown.

Unknown.

Stauridia producta.

ANTHOMEDUS.E — SARSIA.

51

with long, tubular manubrium with the gonad confined to a short length near its free outer
end. He states that the Eximia group are represented by such forms as Sarsia eximia,
brachygaster, flarnmea, barentst, prolijera, angulata, and apiculata. The Tubulosa group are
represented by S. tubulosa, Jensa, decipiens, litorea, pulchella, frutescens, mirabilts, reticulata,
princeps, rosaria, etc.

The genus Plotocnide Wagner, 1885, is defined by Hartlaub, 1907, as a Sarsia-Kks
medusa with nettle-cells upon the exumbrella. The gonad surrounds the manubrium from
the base downward. No ocelli. Hydroid unknown. It appears to me that unless it be
proven that the hydroid is different from Syncoryne, this genus should be merged with Sarsia.

Weismann, 1881 (Zool. Anzeiger, Bd. 4, p. 61), shows that the circulation of fluids
within the gastrovascular cavity of Coryne pusilla is aided by the rhythmical contraction of
the walls of the gonophore. The systole and diastole are not always of equal duration, but
each ranges from 60 to 75 seconds. Thus the circulation in certain hydroids may be aided
by periodic peristaltic contractions as well as by the movement of cilia.

Annandale, 1907, Journal and Proc. Asiatic Society of Bengal, vol. 3, finds Syncoryne
filarnentata, sp. nov., developing free medusae and growing in brackish pools of one-third
the salinity of sea water at Port Canning, Lower Bengal. The mature medusa is unknown.

Tabular Description of the Medusa of Sarsia. — Continued.

S. rosaria= Coryne
rosaria A. Agassiz,
1865.

S. minima von
Lendenfeld.

S. brachygaster
Gronberg,
1898.

S. angulata
Mayer.

S. gracilis
Browne, 1902.

S. princeps
Haeckel, 1879.

Shape and

Oval, with small

Ovate. 3 high,

Three-fourths- Half-egg-shap-

Cylindrical,

Conical with thin

size of bell

apical projection.

2.5 wide.

egg-shaped, ed. 3 High,

thick-walled.

walls. 281040

in mm.

15 to 30 high, 10

widest above 2.8 wide. 4-

5 high, 3 wide.

high, 15 to 30

to 15 wide.

the middle. 15 sided in con-

wide.

to 18 high, 8 traction.

to 10 wide.

Character of

Basal bulbs large;

Tentacles about

Basal bulbs

Basal bulbs

Ocelli on basal

Basal bulbs

4 tentacles.

each with abaxial

2.5 r long.

large with a large, with well-

bulbs. Each

elongate, con-

Length in

ocellus. Tentacles

small abaxial developed ab-

tentacle 3 r

ical, and Banked

terms of

3 to 4 r long.

ocellus. Ten-

axial ocellus.

long, termina-

by pad-like clus-

bell-radius

Basal bulbs

tacles 3 to 4

Tentacles z r

ting in a ni-nia-

ters of nemato-

(r).

flanked by pads

rlong.

long-

tocyst-knob.

cysts. Each bulb

of nettle-cells.

bears small ab-

axial ocellus.

Tentacles 8 r

long.

Shape and

Spindle-shaped,

As in S. rosaria.

As in S. eiimia.

Shaped as in S.

Two-thirds as

Tubular, cylin-

length of

only 1.510 2 r long.

rosaria. r to

long as the

drical, with a

manubrium

one-twelfth r

depth of the

round mouth-

in terms of

long.

bell-cavity.

opening 4 r

bell-radius

long.

(r).

Gonads.

Encircles stomach,

As in S. rosaria.

As in S. eximia.

As in S. rosaria.

CO

As in S. rosaria.

leaving both ends

free. No medusa-

buds.

Color.

Entoderm of stom-

Entoderm

Entoderm of

Entoderm of

(0

Entoderm of ten-

ach ranges from

brown.

stomach and stomach and

tacle-bulbs and

yellow to pink or

tentacle-bulbs tentacle-bulbs

manubrium pur-

reddish-brown to

orange-red. robin-egg

ple. Ocelli

purple. Tentacle-

Ocelli black. blue; never

black.

bulbs yellowish-

red. Ocelli

brown to red.

deep-brown.

Where found.

Pacific coast of

Port Jackson,

Spitzbergen and Bahamas to

Stanlev Harbor,

Greenland and

North America,

New South

Greenland, in Tortugas,

Falkland Is-

Spitzbergen, in

southern Califor-

Wales, Aus-

summer. Florida. Win-

lands.

summer.

nia to Puget

tralia.

ter to mid-

•

Sound .

summer.

Hydroid.

Syncoryne rosaria

Syncoryne min-

Unknown.

Unknown.

(? 0

Unknown.

A. Agassiz and

ima.

Syncoryne sar-

Fewkes.

sii Hartlaub,

1905(0 1=

this medusa a

young Slab-

beria ?

52

MEDUS.E OF THE WORLD.

Sarsia tubulosa Lesson.

Syncorvna sarsii (hydroid), LOVEN, 1835, K. Vet. Acad. Handl. for Ar., p. 275, plate 8, figs. 7-10.

Oceania tubulosa (medusa), SARS, M., 1835, Beskriv og Jagtt., p. 25, plate 5, fig. 1 1 . Also: Syncorine sarsii, 1846, Fauna littor.
Noveg., part. 2, p. 2, tab. i, figs. 1-6.

Syncoryne sarsii, LOVEN, 1837, Archiv. fur Naturgesch., Jahrg. 3, p. 321, taf. 6, fig. 25. — WEISMANN, 1883, Entsteh. Sexuabellen
bei Hydromedusen, Jena, pp. 56, 216. — GARSTANG, 1894, Journal Oxford Club, vol. 2, No. 30, p. 7. — CITRON, 1902,
Archiv. Naturges., Jahrg. 68, pp. I, 26, taf. i, 2 (sensory cells of tentacles). — HINCKS, 1868, British Hydroid Zoophytes,
p. 52, plate 7, fig. 3. — ALLMAN, 1872, Monog. Tubularian Hydroids, p. 275.

Syncoryne sarsi and Sarsia tubulosa, BEDOT, 1905, Revue Suisse de Zool., tome 13, pp. 120, 147 (all literature 1835-1850).

Sarsia tubulosa, LESSON, 1843, Hist. Zooph. Acalephes, p. 333. — SCHI'LZE, 1873, Ueber den Bau von Syncoryne Sarsii, p. 14,
taf. 3.— ROMANES, 1885, Jellyfish, Star-fish, and Sea Urchins, etc., International Scientific Series, vol. 49 (reactions to
stimuli). — LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 212. — HARTLALIB, 1907, Nordisches Plankton, Nr. 12, p. 19, figs. 10-16. —
BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 9; 1895, Proc. and Trans. Liverpool Biol. Soc., vol. 9, p. 246.

Sarsia tubulosa, SARS, i$$$=Sarsia macrorhyncha, BUSCH, 1851; BROWNE, E. T., 1905, Proc. Royal Soc. Edinburgh, vol. 25,
p. 758.

Syncoryne gravata, HINCKS, 1868, Hist. British Hydroid Zooph., p. 53.

Sarsia tubuhsa+S. macrorhyncha, HAECKEL, 1879, Syst. der Medusen, pp. 16, 19.

Sarsia tubulosa + S. pulchella, FORBES, 1848, British Naked-eyed Medusa-, pp. 55, 57, plate 6, figs. ^, 3.

Corynt pusilla, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 340.

(!) Syncoryne Sarsii, HARTLAUB, 1905, Zoolog. Jahrbiichern, Suppl. 6, p. 525, fig. F (hydroid from Terra del Fuego).

Sarsia macrorhyncha, BUSCH, 1851, Beobach. wirbell. Seeth., p. lo, taf. 3, figs. 7-10; taf. 4, figs. I, 2.

FIG. 13. — "Sarsia litorea," after Hartlaub, in Nordisches Plankton.

FIG. 14. — "Sarsia decipiens," after Hartlaub, in Nordisches Plankton.

FIG. 15. — "Syncoryne gravata," after Hincks, in British Hydroid Zoophytes — probably hydroid of Sarsia

T^l- ' . _J ' r 1 _£T^l__ TT 1' t_ . - _1 ' ^1_ - ! L..^ :^ J' _

mirabilis

This medusa is found off the English coast early in the spring, but it disappears before
August. It is evidently an Arctic species, is abundant off the northern coasts of Europe,
and is found off Iceland. It is very closely related to, if not identical with, the American
S. mirabilis, but may possibly be distinguished by its more slender and higher bell, its very
long manubrium, and its somewhat shorter tentacles. All of these characters are, however,
very variable in Sarsia mirabilis, and I have become convinced that the American and
European forms are at most only varieties, one of the other. For details see the tabular
description of medusae of Sarsia.

Romanes, 1885, carried out many interesting physiological experiments upon this
species, and showed that its ocelli are organs for the perception of light. The medusa is
sensitive only to rays between the red and violet, and is strongly attracted by the light. The
smallest part of the bell-margin is capable of initiating and maintaining the rhythm of the
bell, but if the margin be entirely removed all pulsations of the bell instantly cease, while
the cut-off margin continues to pulsate. Stimulation of the subumbrella of the bell causes
contraction of the manubrium (proboscis), and indeed the bell, when deprived of its margin,
still responds by contractions to all sorts of stimuli, chemical, thermal, electrical, or mechan-
ical, although sustained rhythmical pulsation is never resumed.

The hydroid is Syncoryne sarsii, which is common in shallow water along the coasts from
England to Norway. The polypites are spindle-shaped and elongate and have about 12 to

PLATE 4.

Fig. i. Sarsta mirabilts. Abnormal medusa with a tentacle arising from
the side of the manubrium. Nahant, Massachusetts, May 8,
1897. Metschnikoff, 1870, observed a similar abnormality in
Slabbena catenata.

Fig. 2. Median section of a young medusa of Sarsia mirabilis. The ocellus
on the right-hand tentacle is shown as if depigmented in order to
illustrate its structure. The eye on the left-hand tentacle-bulb is
shown normally pigmented.

Figs. 3 and 4. Sarsia mirabilis var. reticulata, young medusa. Figure 4,
one of the nematocyst-cells from the tentacles of figure 3. Agas-
siz Laboratory, Newport, Rhode Island, June 13, 1895.

Drawn from nature, by the author.

ANTHOMEDUS.E — SARSIA. 53

16 short tentacles which arise at irregular intervals from their sides. Each of these tentacles
ends in a knob-like cluster of nematocysts. The 2 or 3 medusa-buds arise from the sides of
the polypite between the tentacles. The stems of the hydroid are quite smooth, sparingly
branched, and about 12 to 15 mm. high. The stems are translucent, slightly horny in color, and
the polypites are light-red.

I can not determine any well-defined distinctions between S. sarsii and S. mirabtlis,
excepting that in S. sarsn the medusa-buds appear always to arise from near the middle of
the sides of the polypite between the tentacles, whereas in S. mirabilts the medusa-buds arise
from near the base of the polypite at or below the level of the lowest zone of tentacles.

Hartlaub has given excellent figures and descriptions of a number of medusae which are
closely related to Sarsia tubiilosa, if not mere varieties of the latter. These are S. pattersoru
Haddon, S. frutescens Allman, 5. decipiens Hartlaub, S. litorca Hartlaub, and S. pulchella
Forbes. (See Hartlaub, 1907, Nordisches Plankton, Nr. 12, pp. 29, 30, 32, 36; figs. 20-226,
23, 24, 28, 29.) I hesitate to quote these as distinct species, for I have observed the same
or nearly the same variations among individuals in swarms of S. mirabilis at Nahant, Woods
Hole, and Newport on our coast. A statistical study, or better still, a study of the respective
hydroids, is required before we can hope to determine these so-called "species" with certainty.

Garstang, 1894, observes that the hydroid of S. tubulosa gives rise to dimorphic medusae,
as does S. mirabilis on the coast of New England, where early in the spring the medusae
are set free, whereas in May they mature while still attached to the hydroid.

Sarsia tubulosa Lesson, variety Sarsia mirabilis L. Agassiz.

Plate 3, figs. 2, 4, and 5; plate 4, figs. 1 and 2.
LITERATURE RELATING TO THE AMERICAN VARIETY or SARSIA TUBULOSA.

Sarsia mirabilis, AGASSIZ, L., 1849, Mem. Amer. Acad., New Ser., vol. 4, p. 224, plates 4, 5. — STIMPSON, 1853, Marine Invert.
Grand Manan, p. u. — HAECKEL, 1879, Syst. der Medusen, p. 17. — FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard
Coll., vol. 8, p. 141, plate 3, figs. 1 1, 12. — LEVINSEN, 1893, Vid. Meddel. Nat. For. Kjobenhavn (5), Bd. 4, p. 143. — BIR-
ULA, 1896, Annuaire Musee Zool. Imp. Sci. St. Petersbourg, tome I, p. 332. — HARTLAUB, 1907, Nordisches Plankton,
Nr. 12, p. 37, figs. 30-40. — LINKO, 1900, Mem. Acad. Sci. St. Petersbourg, ser. 8, vol. 10, No. 4, p. u, taf. I, figs. 1-12
(structure of ocelli); 1905, Zool. Anzeiger, Bd. 28, p. 212; 1900, Traveaul Soc. Jmperiale des Nat. St. Petersbourg, torn.
29, p. 151.

Sarsia mirabilis (medusa), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 21 1-217; vol. 3, Ibid., plate 18, figs. 15-250.

Sarsia glafialis, MORCH, 1857, Beskriv af Gronland, p. 95.

Corync mirabilis (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 185-21 1, plates 17-19; vol. 3, Jbid., figs. 1-16,
plate 17; figs. 1-14, plate 18; figs. 1-27, plate 19.

Corvnt mirabilis, AGASSIZ, A., 1865, North Amer. Acal., p. 175, figs. 282-285. — CALKINS, 1899, Proc. Boston Soc. Nat. Hist.,
vol. 28, p. 336.

Oceania lubulosa, GOULD, 1841, Invert, of Mass., p. 348.

Syncorvnt mirabilis, ALLMAN, 1871, Monog. Tubul. Hydroids, p. 278. — HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol.
24, p. 30, plate v, fig. i; 1901, Amer. Naturalist, vol. 35, p. 578, fig. 33; 1903, Science, vol. 16, p. 344 (variations). —
HARTLAUB, 1905, Zool. Jahrbuchern, p. 526; Zool. Jahrb., Syst. Abth., 1901, Bd. 14, p. 356. — TORREY, 1902, Univ.
California Publications, vol. I, p. 31. — NUTTING, 1901, Bull. U. S. Fish Commission, vol. 19, pp. 328, 372, figs. 3, 81.

( ?) Syndictyon angulatum (young medusa), MURBACH and SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 168.

(})Sarsi barfnisii (young red-colored medusa), LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 214 (north of Russia).

( ?) Euph\sa tentaculata (abnormal medusa with only 3 well-developed tentacles), LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 214
(Barents Sea, North Russia).

Syncoryne densa, HARTLAUB, 1897, Wissen. Meeresuntersuch. Komm. Meere. Kiel, Helgoland, Neue Folge, Bd. 2, p. 452, taf.
166, figs. 4, u ; taf. i6r, figs. 7, 8 (hydroid forms dense tufts). Also: Sarsia sp., HARTLAUB, 1896, Verhandl. Deutsch.
Zool.Gesell., Leipzig, Bd.6, Vers., p. 182 (medusa; with branched manubria); 1907, Nordisches Plankton, Nr. 12, p. 26,
figs. 17-19, 22a (hydroid and medusa).

The following description is derived from a study of medusae and hydroids obtained
off the southern coast of New England, United States.

Adult medusa. — Bell is half-egg-shaped, about 7 mm. in height and 4 mm. in diameter.
There is no apical projection, and the gelatinous substance is not very thick at the aboral
pole and becomes successively thinner near the margin. There are 4 long, highly contractile
tentacles, one at the base of each .radial-canal. The surface of each tentacle is covered with
prominent nematocyst-cells, which are clustered especially at the outer end of the tentacle.
Each tentacle arises from a well-developed basal bulb which contains a single ectodermal
ocellus upon the outer nerve-ring on the abaxial side of the tentacle-bulb. According to
Linko, 1900, the ocellus is composed of a cup-shaped invagmation of densely pigmented
ectodermal cells between which there are spindle-shaped bipolar nerve-cells. The ento-

54

MEDUSAE OF THE WORLD.

dermal core of the tentacle is hollow, and its lumen is continuous with the gastrovascular
system of the medusa. The ectodermal cells of the tentacle-bulb are very thick and are
probably nervous in function. The velum is well developed, being wide and thin. There
are 4 straight, slender radial-canals, and a simple, narrow, circular tube. A short, blindly-
ending tube extends upward from the base of the stomach into the gelatinous substance of the
bell. The entodermal cells of this small projection are several layers thick. The manubnum
is long and extends far beyond the velar opening. Its proximal part is slender and tubular,
but in its outer parts it is much distended by the genital products and consists of a long,
wide, cylindrical tube. The line of demarcation between the narrow and the wide part of
the manubrium is very sharp. The mouth is a simple opening at the extremity of a short
flask-shaped proboscis, and the lips are studded with nematocysts. The mature genital pro-
ducts are found in the ectoderm of the distal part of the manubrium. The entoderm of
the manubrium is usually green, but occasionally it is red. The entoderm of the tentacle-
bulbs is either red or green, and in some individuals the entoderm of the bulb is red while
the ectoderm is green.

FIG. 16. — "Syncoryne pulchella."

FIG. 17. — "Syncoryne frutescens."

The above figures are after Allman, in Ray Society, 1871-72.

Hydroid and young medusa. — The hydroid stock is Syncoryne mirabilis. The stems
are attached by a creeping stolon. They are about 15 millimeters in height and branch
profusely. The main stems and also the side branches terminate each in a single polypite.
The stems are incased in an unannulated chitinous perisarc, which terminates sharply at
the bases of the polypites. Each polypite is fusiform and has about 12 to 18 tentacles which
arise in 3 or more indefinite whorls from the sides of the polypite. These tentacles are not
long, but are quite contractile. Each terminates in a knob-shaped cluster of nematocyst-
cells. The mouth of the polypite is a simple round opening situated at the extremity of a
conical proboscis. Medusa-buds are developed upon the sides of the polypites immediately
below the tentacles, near the lower base of the polypite. Each polypite bears I to 4
medusa-buds in various stages of development. In Massachusetts Bay the breeding season
begins early in March and lasts until the end of May. During March the medusa-buds

ANTHOMEDUS.E — SARSIA.

55

develop 4 long tentacles, and are set free in an immature state, but during the last half of
the breeding season they fail to develop tentacles or give rise to mere short lashes upon their
basal bulbs; and they become sexually mature while attached to the hydroid, the manubrium
of each bud being greatly distended with the genital products. This observation was first
made by L. Agassiz, 1862 (pp. 189, 203), and has been confirmed by us in hydroid stocks
obtained in Swallow's Cave, Nahant, Massachusetts. Plate 3, fig. 5, is derived from one
of these sexually mature medusa-buds found upon a hydroid on May 8, 1897. It will be
observed that the manubrium of the bud is distended with sperm, while the tentacles are not
developed. In this connection it is interesting to observe that Pennaria and Podocoryne
carnca sometimes give rise to medusae which are sexually mature at their time of liberation,
while in other stocks of the same species the medusae are set free in an immature condition.
Garstang, 1894, observes the same phenomenon in the European S. sarsii.

This hydroid of our Sarsia is very abundant, from March until May, in Massachusetts
Bay, where it appears to grow equally well both in pure sea-water and in the brackish mouths
of rivers. The medusae appear in great numbers on the southern coast of New England

FIG. 18. — Hydroid and medusa of "Sarsia Jensa," after Hartlaub, in Nordisches Plankton.

between February and April. They become rare during May, and are not seen during the
summer months. The hydroid extends northward to the Greenland coast, but has not been
recorded from Beaufort, North Carolina, or farther south. Linko and Birula, 1896, found
it in the White Sea, and Linko, 1905, records it from the eastern parts of Barents Sea between
Kanin and Kolgujew Islands. Calkins, Torrey, and Hartlaub have found this medusa
along the Pacific coast of America as far south as Chile. I believe Syncoryne densa Hartlaub
from Helgoland to be an environmental form of S mirabilis.

We have observed an abnormal medusa of Sarsia mirabilis in which a single well-
developed tentacle arose from the side of the manubrium at the point of juncture of the long
tubular basal region and the gemmiferous part of the manubrium. (See plate 4, fig. i). This
abnormal tentacle was studded with clusters of nematocyst-cells. It lacked a basal bulb
and had no ocellus. Asexual budding of medusa- from the walls of the manubrium is not
known in Sarsia mirabilis. Medusae of Sarsia with branched manubria are described by
Hartlaub, 1896, 1907.

Professor Hartlaub finds that in Sarsiti mirabilis the stomach is confined to the distal
end of the manubrium and the gonad is confined to the mid-region of the manubrium above
the stomach. Both the proximal and distal ends of the manubrium lack the gonad. On the
other hand, in S. brachygaster and S. cximia there is no differentiated stomach-region, and
the gonad may extend over the whole, or nearly the whole, length of the rnanubrium.

56

MEDUSAE OF THE WORLD.

The chief and possibly only well-marked point of difference between the forms S. mira-
bilis and S. tubulosa is that in the hydroid of S, mirabilis the medusa-buds arise from near
the base of the polypite, whereas in the hydroid of 5. tubulosa they arise from points higher
up on the sides of the polypite, between the tentacles.

20.

FIG. 19. — Sarsia eximia, after Hartlaub, in Nordischcs Plankton.

FIG. 20. — Syncoryne eximia, after Allman, in Ray Society, 1871-72. Hydroid'and young medusa.

PLATE 5.

Fig. I. Sarsia angulata. Nassau Harbor, New Providence Island, Bahamas,
July, 1903.

Fig. 2. Corynitis agassizii. Charleston Harbor, South Carolina, September,
1897.

Fig. 3. Ectopleura minerva. Tortugas, Florida.

Fig. 4. Ectopleura dumortieri, young medusa. Agassiz Laboratory, New-
port, Rhode Island, June, 1893.

Fig. 5. Ectopleura dumortieri, mature male. Agassiz Laboratory, Rhode
Island, July, 1896.

Fig. 6. Sarsia mirabilis var. reticulata. Nahant, Massachusetts, March 25,
1897.

Drawn from life, by the author.

PLATE 5

AXTHI i.MKDTS.E SARSIA. 57

Sarsia mirabilis var. reticulata.
Plate 4, figs. 3 and 4; plate 5, fig. 6.

Syndictyon reticulatum, A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 340. — AGASSIZ, A., 1865, North Amer.

Acal., p. 177, figs. 290-300.

Syncor\ne reticulata, ALLMAN, 1871, Monog. Tubul. Hydroids, p. 283.
Syndictvon reticulatum, HAECKF.L, 1879, Syst. der Medusen, p. 21.

( ?) Syndictyon raiculalum, MAAS, 1893, Ergeb. der Plankton-Expedition, Bd. 2, K. c., p. 67.
(tySarsia turrreula, MrCRADV, 1857, Gymn. Charleston Harbor, p. 36, plate 8, figs. 6-8.
S\ncor\ne reticulata, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 30.
Sarsia reliculata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 45, figs. 41-43.
(?) Sarsia pulchella, AI.I.MAN, 1871, see Hartlaub, 1907, Nordisches Plankton, Nr. 12, p. 34, fig. 27 (abnormal twin medusa:,

fig. on p. 109). — SPAGNOLINI, 1876, Catalogo Acalefi Mediterranco, p. 18, tav. 2, figs. i. 2.

Adult medusa. — Bell ellipsoidal in shape, being about 4 mm. in height and 3.5 mm. in
diameter. No apical projection. Gelatinous substance quite thick at the aboral pole, but
thin at the bell-margin. There are 4 long, highly contractile tentacles, I at the base of each
radial-canal. Surfaces of these tentacles covered with prominent nematocyst capsules. Basal
bulbs of tentacles well developed and each one contains an ectodermal ocellus upon its outer
side. Velum wide and thin. There are 4 straight, narrow radial-canals, and a slender circular
vessel. Manubrium short and club-shaped, and does not extend far beyond the velar opening.
Mouth a simple, round opening. Genital products developed along the greater part of the
length of the manubrium. The entoderm of the manubrium and tentacle-bulbs is brick-red.

Hydroid and young medusa. — Smaller than Coryne mirnliilis, being not more than 3 mm.
in height. Stems slender and hardly ever branch, excepting in old specimens, which some-
times give rise to a single branch near the base of the stem. Polypites large and club-shaped,
and having several whorls composed of 8 to 10 short tentacles. The medusae develop
among the tentacles near the proximal base of the polypite. When set free the young medusa
is remarkably large, being about 1.5 mm. in diameter. The bell is covered \\irh reticulated
clusters of nematocyst-cells (plate 4, figs. 3, 4) which are especially numerous near the bell-
margin above the circular canal. Some of these nematocyst-cells are large and round, while
others are narrow and long. The tentacles are thickly covered with helically arranged clusters
of nematocyst-cells. These cells (plate 4, fig. 4) are ellipsoidal in shape and are mounted
upon a short basal pedicel. Each cell gives rise to a long, sharp-pointed, sensitive hair. The
nematocyst thread lies coiled in a helix within the cavity of the cell. As the medusa becomes
mature the reticulated nematocysts disappear from the surface of the exumbrella and the
nematocysts upon the tentacles become less prominent. This medusa is found upon the Ne\\
England coast from April until June. It is distinguished from Sm-sm mirabilis only by its
nematocyst-covered tentacles and exumbrella and its small hydroid. It appears also to be
constantly brick-red, while 5. mirabilis is highly variable in color. It is often impossible
to distinguish mature medusae of S. reticulata from those of .V. mirabilis. It is possible that
the S. pulchella of Spagnolini, 1876, from Naples, Italy, is identical with .S'. r, n, uLita.

Sarsia eximia Boehm.

Coryne eximia, ALLMAN, 1859, Annals and Mag. Nat. Hist., ser. 3, vol. 4, p. 141; 1864, Ibid., vol. 13, p. 357.

Syncoryne eximia, HINCKS, 1868, Hist. British Hydroid Zooph., p. co, plate 9, fig. 2. — ALLMAN, 1871, Monog. Tubul. Hydroids,

p. 282, plate 5.
Sarsia eximia, BOEHM, 1878, Jena. Zeitschrift fur Naturw., Bd. 12, p. 191, taf. 6, figs. 7-26; taf. 7, figs. 1-6. — (In part), HAEC-

KEL, 1879, Syst. der Mcdusen, p. 17. — HARTLAIPII, 1894, \Vissen. Meercsuntcrsuch. Komini^. Mo-rc. Kid, lldi;ii]and,

Ser. 2, Bd. i, p. 187.

Svncorvne eximia (hvdroid). NUTTINC;, 1901, Proc. Washington Acail. Sa., vol. 3, p. 166, plate 14, figs. 3, 4.
Sarsia eximia, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 21;, p. 756 (hydroid and medusa). — BROCB, 1905, Bergcns

Museums Aarbog, No. 11, p. 4.

Sarsia tximia=S. bretonica, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 8, figs, i, 2a, 26 (full list of recent literature).
Syncorync eximia, BROWNE, 1907, Journal Marine Biol. Association, vol. 8, p. 37 (growth of the hydroid).

This form is found off the coasts of Great Britain, Helgoland, Shetland Islands, Norway,
and Juneau, Alaska (Nutting). For details of the medusa, see tabular description of the
medusas of Sarsia.

The hydroid is about 30 mm. high, forming a bush-like cluster of profusely branched
stems; the branches are short and simple and arise very irregularly from the main stems,
and are usually faintly ringed at their points of origin. The main stem is usually unringed,

58

MEDUSJE OF THE WORLD.

except at its base, and is quite smooth. The polypites are very elongate, spindle-shaped,
and have about 1 6 to 24 short, knobbed tentacles arranged in 4 to 6 somewhat irregular
verticils. The medusa-buds arise singly upon short peduncles near the bases of the tentacles
of the lower verticils. The entoderm is red to reddish-brown and the stems are yellow.

Browne found that confinement in an aquarium under somewhat unnatural conditions
caused the hydroid to grow rapidly and to form stolons, these being developed from branches

which touched the glass sides of the aquarium.
He also discovered that the medusa becomes
mature in from J to 10 days after being set free
from the hydroid. Later, in 1907, Browne found
that one of these hydroids placed in a glass tube
with a constant current of water passing through
it grew in length from 14 to 77 mm. in the course
of 9 days, and developed branches having a total
length of 500 mm. The hydroid was fed upon
copepods.

Sarsia radiata von Lendenfeld.

Sarsia radiata, VON LENDENFELD, 1884, Zool. Anzeiger, Jahrg. 7,
p. 584; 1884, Proc. Linnean Soc. New South Wales, vol.
9, pp. 583, 635; plate 20, figs. 31, 32; plate 30, figs. 1-4.

Medusa. — Bell semiovate, slightly higher
than broad, 3 mm. high, 2.5 mm. wide. 4 ten-
tacles, each about 1.5 times as long as bell-height,
and with large bulbs about half as wide as the
manubrium. Ocelli (?) Velum wide. 4 straight
radial-canals. Manubrium cylindrical, half as
long as the bell-height. The gonad incases the
sides of the manubrium from the inner apex of
the bell-cavity to near the mouth. No medusa-
buds. Entoderm of manubrium and tentacle-
bulbs deep brown. Other parts colorless.

H \droid. — The hydrocauli arise from a
creeping hydrorhiza which anastomoses in a
very open network. The perisarc terminates
with an oblique elliptical margin at the base
of each hydranth, and the hydranth is provided
with a muscle at this point which enables it to
bend downward and "shut up" as if it were the
blade of a penknife. The hydranths are spindle-
shaped, narrow, and elongate; and are, includ-
ing their hydrocauli, 3 to 5 mm. high. They have 6 to 8 verticils, each of 4 tentacles, situated
in 4 meridional lines, 90° apart. These tentacles are all knobbed at their ends. The hydranths
which produce medusae are shorter than the sterile polypites. The medusae bud out from the
lower half of the polypite between the tentacles. The entoderm is intensely brown in color,
and the perisarc is bright brownish-yellow. Other parts colorless. Found on the coast of
New South Wales, Australia. The medusae are produced in April and May.

Sarsia conica.
Codonium eonicum, HAECKEI., 1880, Syst. der Medusen, p. 634.

Bell barrel-shaped with conical apex one-third as long as the sides of bell. 12 mm. high,
4 mm. wide. 4 tentacles longer than bell-height, and with small oval basal bulbs. The
manubrium is half as long as the depth of the bell-cavity. The stomach is subspherical and
swollen by the encircling gonad. The mouth is at the end of a short, cylindrical throat-tube
which is free of gonads. Color ( ?) There is a long axial canal above the stomach. Indian
Ocean. Briefly described, without figures, by Haeckel.

FIG. 21. — Sarsia eximia, from life, by the author. Mouse-
hole, Cornwall, England, Nov. 14, 1907.

A\THi>MKI>rs.K--SAi;-.I \. .V.I

Sarsia rosaria Haeckel.

Coryne rosaria, AGASSIZ, L., 1862, Cent. Xat. Hist. U. S., vol. 4, p. 340. — A- \ i/. \.. i Sf>;, Virrh Arm-r. Aca!., p. 176, fig 289.

Sarsia rosaria, HAECKEL, 1879, Syst. Jer Mi .i'i CDj p. 18.

Santa rosaria= Syncoryne rosaria, FEWKES, iSSij. Am. r. Vituralist, vol. 23, p. 597, plate 25, fig. 7; tcit-figs. 8, 9 (hydroid?).

Syncoryne occidentals, FEWKKS, 1889, Bull. Essei Inst., Salem, vol. 21, No. 7, p. 99, plate 3, figs. 2, 3.

( ?) Syn diclyon angulatum, Mt RUAI H and SIIK^RV.R, 1903, Proc. Zoo]. Soc. London, vol. 2, p. 168.

Codonium apiculum, MURBACH and SHEARER, 190}, Proc. Zool. Soc. London, vol. 2, p. 165, plate 17, fig. i; plate 22, figs. 4, 5;

1902, Annals and Ma^. Nat. Hist., scr. 7, vol. 9, p. 72.
Sarsia apicula+S. rosaria, HARTLAI'B, 1907, Nordischcs Plankton, Nr. 12, pp. 17, 50, figs. 9, 45.

Bell 15 to 30 mm. high, 10 to 15 mm. wide, with fairly thick walls and small apical
projection. 4 equally developed, radially placed tentacles 1 .5 to 2 times as long as bell-height.
The basal bulbs of these tentacles are large and are flanked on either side by a large nemato-
cyst-pad. Each tentacle-bulb bears an abaxial ocellus. There are 4 slender, straight-edged
radial-canals and a narrow ring-canal. The velum is well developed. Manubrium short and
spindle-shaped, and mouth about at the level of the velar opening. There is a short axial
canal above the stomach. The gonad encircles the stomach, leaving both ends tree. N<>
medusa-buds. The colors are quite variable as in other species ot Sarsia. The tentacle-
bulbs range from yellow through red to brownish-red, and the stomach is yellow, pink, or
reddish-violet to purple.

This is the most abundant Sarsia along the Pacific coast of the United States. It occurs
in great swarms in San Francisco Harbor in spring; and in Victoria Harbor, Puget Sound,
in July.

The hydroid is Syncoryne rosaria found by A. Agassiz and Fewkes in shallow water
attached to piles of wharves. Each tentacle terminates in a knob, and the hydroid is a true
Syncoryne.

Sarsia minima von Lendenfeld.
Sarsia minima, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, pp. 584, 915, plate 21, figs. 34, 35.

Bell of medusa 3 mm. high and 2.5 mm. wide with "a long manubnum like the northern
Sarsia;." It is therefore readily distinguished from Sarsia radiata, which has a short manu-
brium. S. minima has a spindle-shaped, nearly cylindrical manubnum which extends tor
about half its length beyond the velar opening. The 4 marginal tentacles are somewhat
longer than the bell-height and are covered with rings of nematocysts. The entoderm ot
the stomach is pale brown, other parts colorless.

Hydroid. — The stems are 2 to 3 mm. high and arise from a creeping, slightly branched,
non-anastomosing hydrorhiza. The perisarc which invests the hydrorhiza and hydrocauli is
irregularly annulated or wavy throughout, and terminates at the bases of the hydranths in
a transverse margin. The hydranths are slender, spindle-shaped, 0.6 to 0.8 mm. long, and
with 8 to 12 irregularly scattered tentacles, all of which are knobbed at their ends. When
they produce buds they become stouter, and are so thickly covered by the budding medusae
"that nothing of their bodies remains visible." The perisarc is reddish-brown.

This hydroid is found at Port Jackson, New South Wales, Australia, overgrowing Obclla
geniculata, on buoys and submerged ropes. The medusae are produced in April and May.
Von Lendenfeld did not obtain any mature medusae.

Sarsia brachygaster Grbnberg.

Sarsia brachygaster, GRONBERO, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 1 1, p. 459, taf. 27, figs. 3, 4. — HARTLAI n, 1907, N'or-
disches Plankton, Nr. 12, p. n, fig. 3.

Bell 15 to 18 mm. high and 8 to 10 mm. wide and three-fourths-egg-shaped, the greatest
breadth being above the middle. 4 radially situated tentacles, each being about twice as
long as the bell-height. Basal bulbs of these tentacles well developed, and a single MI\
small ocellus upon the outer side of each bulb. Manubrium cylindrical, and two-thirds
as long as height of bell-cavity. Mouth situated at extremity of a shoit cylindrical neck.
The gonad is tubular and surrounds the stomach. No medusa-buds. The manubrium,
gonads, tentacle-bulbs, and tentacles are orange-red. The ocelli are black.

60

MEDUS.E OF THE WORLD.

Gronberg found this species at Spitzbergen in summer, and he also identified it among
a collection of medusae from Jakobshavn, Greenland, where it appears to be rarer than at
Spitzbergen.

Sarsia angulata Hartlaub.
Plate 5, fig. i; plate 6, fig. 3.

Syndictyon angulatum, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College., vol. 37, p. 5, figs. 6-8, plate 3; 1904,

Memoirs Nat. Sci. Brooklyn Institute Museum, vol. I, No. I, p. 7, plate I, fig. 6.
N on Syriilict\on tiHgtihitum, MURBACH and SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 168.
Sarsia angulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 16.

Bell i mm. high; half-egg-shaped, with moderately thick walls. Becomes almost square
in cross-section when contracted. There are 4 slender tentacles with fairly thick spindle-
shaped ends. These tentacles are each about as long as bell-height and their distal halves are
tapering and are armed with nematocysts. The basal bulbs of the tentacles are not very large,
and each one bears an ocellus formed by a cup-like invagmation of ectodermal cells. The velum
is large, and the radial-canals and circular vessel are of fine caliber. The manubrium is
spindle-shaped with a narrow tubular o?sophagus and without an aboral projection. It is
about two-thirds as long as the height of the bell-cavity. The gonad encircles it, extending
from the base to near the mouth, leaving the throat-tube free. The entoderm of the tentacle-
bulbs and manubrium is robin-egg blue, while the ocelli are deep-brown, almost black. All

other parts are hyaline. This medusa is abundant
in the Tongue of the Ocean, Bahama Islands, in June
and July, and was found at Turks Island in January.
It is rare at Tortugas, Florida.

Sarsia gracilis Browne.

Sarsia gracilis, BROWNE, 1902, Annals Mag. Nat. Hist., ser. 7, vol. 9, p. 275.
(? ?) S\ncoryne sarsii (hydroid), HARTLAUB, 1905, Zoolog. Jahrbiichern,
Suppl. 6, p. 525, fig. F.

Bell 5 mm. high, 3 mm. wide; cylindrical, with
moderately thick walls and quadrangular margin.
4 tentacles, about as long as the bell-height, and end-
ing each in a large knob containing nematocysts. An
ocellus on the basal bulb of each tentacle. Manu-
brium about two-thirds as long as the depth of the
umbrella cavity. Color ( ?) Gonads ( ?) Found at
Stanley Harbor, Falkland Islands, by Vallentin, and
briefly described without figures by Browne. The
hydroid, Syncoryne sarsn, described by Hartlaub from
southern Terra del Fuego, may be the stock ot this
medusa.

The medusa may be a young Slabberia ( ?)

Sarsia princeps Haeckel.

Codonium princeps, HAECKEL, 1879, Syst.derMedusen,p.l3,taf.l,figs. 1,2.

Sarsia princtps, HAECKEL, 1879, Ibid., p. 655. — LINKO, 1905, Zool. An-
zeiger, Bd. 28, p. 212. — BROWNE, i9O3,Bergens Museums Aarbog,
No. 4, p. 8, plate i, fig. i; plate 3, fig. 4. — HARTLAUB, 1907, Nor-
disches Plankton, Nr. 12, p. 47, fig. 44.

Codonium princeps, LEVINSEN, 1893, Vid. Meddel. Nat. For. Kjobenhavn
(5), Bd. 4, p. 143. — GRONBERG, 1898, Zoolog. Jahrb., Abth. Syst.,
Bd. n, p. 458, taf. zj, figs, i, 2.

Bell thin-walled and conical, about 25 to 40 mm.
high, and 15 mm. wide. There is a short, conical,
apical projection. There are 4 tentacles with long
conical basal bulbs. The shafts of these tentacles

1-iG. 22. — barsia princeps, alter Hartlaub, in

Nordisches Plankton. are very contractile, three or four times as long as

PLATE 6.

Figs, i and i'. Hydroid of Ectopleura Jumortieri. Dredged from a depth

of 5 fathoms in Newport Harbor, Rhode Island, August, 1896.

Figure I, enlarged view; figure i', natural size.
Fig. 2. Medusa of Ectopleura dumortteri. Agassiz Laboratory, Newport,

Rhode Island, July, 1892.

Fig. 3. Sarsia angulata. Turks Islands, Bahamas, January 2O, 1893.
Fig. 4. Protiara formosa, male. Nassau Harbor, Bahamas, June, 1903.
Fig. 5. Protiara formosa, female. Tortugas, Florida, June IO, 1897.
Fig. 6. Protiara formosa, young medusa. Turks Islands, Bahamas, January

20, 1893.
Fig. 7. Zanclea gemmosa. Manubrium of the medusa shown in figure 5,

plate 7.

Drawn from life, by the author.

PLATE 6

ANTHOMEDUS.fi — SARSIA.

61

the bell-height, and covered with alternately arranged, wart-like clusters of nematocysts. A
small ocellus is situated on the outer surface of each tentacle-bulb near the bell-margin. There
are also 2 globular swellings, one on either side of each tentacle-bulb adjacent to bell-margin.
The velum is narrow. There are 4 narrow, ragged-edged radial-canals and a slender circular
canal. A short axial canal extends upward from the stomach-cavity into the gelatinous
substance of the apical projection of the bell. The manubrium is long and cylindrical, and
extends for about one-third of its length beyond the velar opening. The mouth is a simple
opening without prominent lips. A single, short, tubular gonad is developed upon the sides
of the manubrium. The manubrium, gonads, and tentacle-bulbs are purple. The ocelli
are black.

This species is found off the coasts of Greenland and Spitzbergen, where it appears to
be common. Gronberg, 1898, found it to be abundant at Spitzbergen, and Linko, 1905,
found it to be common in Barents Sea, north of Lapland, Russia. It is the largest known
Sarsia. Gronberg's description is based upon the study of living medusae. Hartlaub also
gives an excellent figure of the medusa, which we reproduce.

FIG. 23. — Sarsia prolifera, from life, by the author. Mouse-
hole, Cornwall, England, Nov. 8, 1907.
FIG. 24. — Sarsia codonophora, after Haeckcl, 1879.

Sarsia prolifera Forbes.

Sarsia proliftra, FORBES, 1848, British Naked-eyed Mi-ilus.r, p. 59, plate 7, fig. 3. — BUSCH, 1851, Beobacht. wirbeilos. Seeth.,
p. i, taf. i, figs. 1-6. — BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 147 (all literature to 1850). — HARTLAUB, 1907,
Nordisches Plankton, Nr. 12, p. 15, figs. 7, 8.

Syncorync prolifera, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 83, fig. 38.

Codonium codonophorum+ Sarsia prolifer, HAECKEL, 1879, Syst. der Medusen, pp. 14, 18, taf. I, fig. 3.

Sarsia codonophora, HAECKEL, 1880, Ibid., p. 655.

62 MEDUS.'E OF THE WORLD.

In 1848 Forbes described this medusa from Penzance Bay, southern England, where
it was abundant in August, 1846, and in 1879, and where I found it in November, 1907.
Haeckel describes that which may prove to be the same species from the Mediterranean. The
points of difference between the two medusae will be brought out in the following description.

The bell is bell-shaped; in Forbes's medusa (fig. 23) 3 mm. wide and 4 high; in Haeckel's,
(fig. 24), 8 mm. wide and 8 mm. high. Forbes's medusa lacks an apical projection, while
Haeckel's has a short, conical, pointed apex. Also, in Forbes's medusa there was no axial
canal above the stomach while in Haeckel's there was a narrow, axial vessel. Moreover,
in Haeckel's medusa the margin of the bell flares outward, while in Forbes's specimen this is
not the case.

It is well known to students of the Anthomedusae that apical projections and axial canals
are exceedingly variable, and may be absent or present in individuals of the same species.
The outward flaring of the bell may be brought about through contraction. Haeckel's medusa
appears to be merely a large, highly-colored specimen of S. prohfera Forbes.

The 4 marginal tentacles are I to 3 times as long as the bell-height. Their basal bulbs
are large and tapering and about as wide as they are long. The tentacles taper gradually
from base to shaft and are hollow. Clusters of I to 3 medusa-buds at a time are developed
upon the tentacle-bulbs immediately below the bell-margin. When set free these medusas
are already producing another asexual generation of medusas upon their tentacle-bulbs.
The 4 radial-canals and ring-canal are straight and very narrow. The manubnum is spindle-
shaped and nearly as long as the bell-height in Haeckel's, but not half so long in Forbes's
medusa. The mouth is at the end of a narrow neck and is a simple, round opening. The
stomach is encircled by the gonad, leaving the base and the throat-tube free.

Entoderm of tentacles and manubnum yellow to orange or sage-green. Each tentacle-
bulb bears a prominent ectodermal brown-colored ocellus.

If it be true that Forbes's and Haeckel's forms are identical the medusa must range from
southern England to the Mediterranean. It appears to be rare and its hydroid is unknown.

Allman, 1871 (p. 83, fig. 38). gives a figure of a medusa which closely resembles Forbes's
Sarsta prohjera.

I found numerous specimens of this medusa off Mousehole, Mounts Bay, Cornwall,
England, early in November, 1907. They accord well with the descriptions of Forbes
and of Busch. The entoderm of the manubrium and tentacle-bulbs was sage-green; the
tips of the tentacles light reddish-brown, and occasionally there was some reddish-brown
pigment in the central entoderm of the stomach. The gonad encircled the stomach, leaving both
ends of the manubrium free.

Sarsia gemmifera Forbes.

Sarsia gemmifera, FORBES, 1848, British Naked-eyed Medusae, p. 57, plate 7, fig. 2. — CHUN, 1895, Bibliotheca Zoologica, Heft
19, pp. 4, 7, taf. I, figs. 5, 6 (law of development of medusa-buds). — BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25,
p. 757. — BROCH, 1905, Bergens Museums Aarbog, No. 1 1, p. 4.

Codonium gemrniferum+ Sarsia sipfionophora, HAECKEL, 1879, Syst. der Metlusen, pp. 15, 20, taf. I, fig. 4.

Codonium gcmmifcrum, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 132 (citation of literature to 1850).

Purena gemrtiifera, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 58, figs. 54-58.

This medusa is tound off the Atlantic coasts ot Europe from Norway southward.
Haeckel's Sarsia "siphonopliora" from the Canary Islands is probably another name for the
same species.

When young the medusa produces medusa-buds upon its manubrium. These arise in
a spiral line, one following the other, down the sides of the manubrium. The oldest of the
primary medusa-buds is nearest the base (uppermost) and the youngest nearest the mouth
of the manubrium.

These primary medusa-buds are attached to the sides of the manubnum by short pedicels,
and secondary and tertiary medusa-buds arise from the sides of these pedicels. Thus when
the oldest (uppermost) original medusa-bud is set free, the secondary bud upon its pedicel
remains attached to the manubrium and continues to develop, and when it in turn is set free
the tertiary bud completes its development; this process takes place with each and every
one of the series of buds.

ANTHOMEDUS.E — SARSIA.

63

This successive freeing of the three series of buds produces various appearances at different
periods of time, and in this manner we may explain the condition observed by Haeckel, 1879,
in Sarsia " siphonop/iora," where the buds appeared in two series, the oldest and largest of
the upper series being at about the middle ot the manubnum, while the oldest ot the lower
series was near the base of the stomach. Forbes, 1848 ( plate 7, fig. 2 e), shows an intermediate
stage in the budding process, and an early stage is shown by Chun, 1895 (fig. 25 A). The
law of succession in development of the medusa-buds is clearly demonstrated by Chun.

Both ectoderm and entoderm of the
manubrium take an equal share in the
formation of the budding medusae.

For details of the character of the
medusa, see tabular description of the
medusae of Sarsia.

Haeckel failed to observe the sec-
ondary and tertiary medusa-buds upon
the pedicels of the budding medusae in
his Sarsia siplionophora, but this is
probably due to an oversight, tor in all
other respects his medusa appears to be
identical with S. gemmiftra P'orbes.

The gonad develops at the distal
end of the manubrium after the budding
process has ceased. Possibly there may
be 2 or more ring-like gonads ? (See
Hartlaub, 1907.) If this be the case
the medusa may be identical with
Dipuretia fertilis Metschnikoff, 1871.

Sarsia hargitti.

Corynt producta, HARGITT, 1902, American Natural-
ist, vol. 36, p. 550, fig. 3.

Syncorynf produtla, HAROITT, 1904, Bulletin Bureau
of Fisheries U. S., vol. 24, p. 30, plate I, fig. I.

A medusa called Sarsia producta is
produced by budding from the hydroid
of StauriJia producta Wright; and as
this name takes precedence over that
of Hargitt's medusa, it is necessary to
rename the American species.

Bell i .5 mm. high, and I mm. wide.
Apex dome-like and rounded, and sides
slightly compressed. Bell-cavity only
about half as deep as the height of the
bell. There are 4 long, equally devel-
oped tentacles with large basal bulbs,
each with a large, ectodermal ocellus.
Velum well developed. The 4 radial-
canals are narrow, straight, and smooth-
edged. The manubrium projects far
beyond the veJar opening, but is cap-
able of great contraction. Its terminal
part is bulb-like, while the basal portion is spindle-shaped and bears a whorl of medusi-
1'orin gonads. The stomach gives rise to a blunt, dome-shaped aboral projection. The
mouth is a simple, round opening. The basal part of the manubrium is orange, and its
free end blue-green. The tentacle-bulbs are orange, edged with delicate green. The ocelli
are black. A single specimen was found by Hargitt at Woods Hole, Massachusetts, on August

FIG. 25. — Sariia gcmmifera, showing law of building. After Chun, in

Bibliotheca Zoologica, Heft 19, taf. I, 1895.
A, early; B, later stages of the process of budding.

64

MEDUSAE OF THE WORLD.

10, 1901. It is distinguished from all other American species of Sarsia by its medusiform
gonads borne upon the manubrium (text-fig. 26). It is not known whether medusae are set
free from the manubrium of the parent medusa.

Sarsia flammea Hartlaub.

Sarsia eximia (in part), HAECKEL, 1879, Syst. der Medusen, p. 17.

Sarsia fammea, LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 212. — HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 12, figs. 4-6.
(literature); 1903, Zool. Centralblatt, p. 22.

Bell high oval, 12 mm. high, 7 mm. wide, walls quite uniform and of moderate thickness.
4 tentacles, with well-developed, simple basal bulbs without ocelli. Tentacle tips slightly
enlarged, club-like. Nematocysts over distal halves of tentacles arranged in prominent, broken
rings. 4 slender, straight radial-canals. Manubrium thick, conical, spindle-shaped; only
two-thirds as long as the depth of the bell-cavity. No axial-canal above the stomach. Gonad
ring-like, encircling the manubrium from base to near the mouth. Stomach and tentacle-
bulbs light fiery-red or orange.

From the Arctic Ocean.

FIG. 26. — Sarsia hargitii, after Hargitt, in Bull. TJ. S. Bureau of Fisheries.

FIG. 27. — Sarsia flammea, after Hartlaub, in Nordisches Plankton. Showing arrangement of nemato-
cyst clusters on tentacles.

Separated from Sarsia eximia and S. brachygaster by its lack of ocelli, and from S.
brachygaster also by its smaller size. The best description is that of Hartlaub, 1907. Hydroid
unknown. It is possible that this may be the medusa of the parasitic hydroid called
Hydnchthys mirus Fewkes.

Genus SARSIA, Subgenus STAURIDIOSARSIA nov. subgen.
Stauridium, HARTLAUB, 1895, Zeit. fiir wissen. Zool., Bd. 61, p. 142; 1907, Nordisches Plankton, Nr. 12, p. 52.

CHARACTERS OF THE SUBGENUS.

Medusa similar to Sarsia, but the hydroid is Stauridia, not Syncoryne.

The name Stauridia should not be applied to these medusae, for it was first used by
Dujardin, 1843, to describe a hydroid which gives rise to a Cladonema medusa. It may be
well to apply the new generic name Stauridiosarsia to hydroids which resemble Stauridia,
but produce Sarsia-\\ke medusae.

ANTHOMKDUS.E — SARSIA.

65

Sarsia (Stauridiosarsia) producta.

A^on Stauridir, DUJARDIN, 1843, Ann. Sci. Xaturelles, scr. 2, torn. 20, p. 370.

Stauridia producta, WRIGHT, 1858, Edinburgh New Philos. Journ., ser. 2, p. 283, plate 7, figs. 6-8.

Stauridium productum, HINCKS, 1868, British Hydroid Zoophytes, p. 68, plate 12, figs, i, I a. — HARTLAUB, 1895, Zcit. fur

wissen. Zool., Bd. 61, p. 142, taf. 7, figs. 1-19; taf. 8, figs. 1-4; taf. 9, figs. I, 2, 6; 1907, Nordisches Plankton, Mr.

12, p. 53, figs. 47-50.

The medusa is a Sarsia, but the hydroid is similar to Stauridia, and differs from Syn-
corync, the hydroid of Sarsia, in that there is a basal circlet of simple, knobless tentacles;
whereas all of the tentacles of Syncoryne terminate in knobs. The tentacles of Stauridia

28.

FIG. 28. — Sarsia producta, after Hartlaub, in Nnrdisches Plankt»n.

FIG. 29. — Medusa; of "Sarsia producla," natural size; after Hartlaub, in \ordischt-s Plankton.

FIG. 30. — "Stauridium productum," after Hincks, in British Hydroid Zoophyte-;. Hydroid and young medusa.

are of two sorts — the simple, short, stiff, tapering, knobless basal circlet, and above them
several circlets of knobbed tentacles which arise from the sides ot the hydranth. In Syncoryne,
on the other hand, we find only knobbed tentacles. Thus two distinct genera of hydroids
produce one and the same genus of medusa. Moreover, Dujardin, 1843, discovered that the
hydroid which he called "StauriJie" produces the medusa Cladonema rnJititutu.

As the name Stauridia is preoccupied by Dujardin to designate the hydroid of Cladonema
it can not be applied to the medusa of Stauridia producta. I therefore call the medusa of
Stauridia producta by the name Sarsia producta, thus indicating its relationships when in
the reproductive stage.

The medusa Sarsia producta, which is produced by the hydroid Stauridia producta, may
be described as follows:

Bell 10 mm. high, 7 mm. wide, three-fourths-egg-shaped, with thick, gelatinous walls.
4 equally developed, radially placed tentacles; 1.5 times as long as the bell-height. Tentacle-

66

MEDUSA OF THE WORLD.

bulbs large, each with an abaxial, ectodermal ocellus. Manubrium cylindrical, with about
one-third of its length extending beyond the velar opening. A short, conical axial canal above
the stomach. Gonad extends from the base to the distal end of the manubnum. No medusa-
buds. Stomach brownish, gonads yellowish-white, tentacle-bulbs and axial canal red.
Helgoland and British coasts.

The hydroid has a thin-branching hydrorhiza, from which a number of club-shaped
hydranths arise singly. These hydranths are each about 2 mm. long, with a proximal circlet
of 4 to 6 short knobless tentacles, and above 2 or 3 more or less irregular circlets of large
knobbed tentacles, usually with 4 tentacles in each circlet. The medusa-buds are produced
from the sides of the hydranth above the basal circlet of tentacles. The hydranths are red.
The hydroid and medusa are described in detail by Hartlaub, 1895.

The English and Helgoland forms differ considerably and they may be regarded as
varieties one of the other (see Hartlaub, 1895, p. 157). The hydrorhiza of the English form
anastomoses, while that of the Helgoland hydroid branches sparingly and does not anasto-
mose. In the English form the
knobbed tentacles are in whorls of
4 each, whereas in the Helgoland
hydroid there areotten 5, occasionally
6, tentacles in each whorl. In the
English form the medusae are said
to arise from the bases of the ten-
tacles, while in the Helgoland form
they arise from the sides of the
polypite between the knobless basal
circlet and the lowest circlet of
knobbed tentacles. Finally the Hel-
goland medusa has a largeaxial canal
above the stomach, and this appears
to be absent in the English medusa.

Genus HYDRICHTHYS Fewkes, 1888.

H\drichthys, FEWKES, 1888, Bull. Mus. Comp.
Zool. at Harvard Coll., vol. 13, p. 224;
1888, Annals and Mag. Nat. Hist., ser. 6,
vol. i, p. 364. — HARGITT, 1904, Bull. U. S.
Bureau of Fisheries, vol. 24, p. 32.

( ?) Plotocnide, HARTLAUB, 1907, Nordisches Plank-
ton, Nr. 12, p. 68.

GENERIC CHARACTERS.

The medusa resembles Sarsia,
but is without ocelli upon the ten-
tacle-bulbs. The mature medusa has
not been determined. The hydroid is
firmly attached to the side of a fish
(Sertola zonata), and is probably
parasitic, and degenerate in many
respects. All of our knowledge of
this remarkable form is derived from
Fewkes, 1888. It differs widely from
all other known forms of Tubulanan
hydroids.

In default of knowledge of the
mature medusa we must remain in
doubt concerning its true place in
our classification. It is possible that
Sarsia flamrnea Hartlaub may be the mature stage of this medusa.

\

FIG. 31. — Young medusas of Hydrichthys rnirus, from Fewkes, in Bull.
Museum Comp. Zool. at Harvard College.

ANTHdMKDUS.E — HYDRICHTHYS.

67

Hydrichthys mirus Fewkes.

Hydrichthys mirus, FEWKES, 1888, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 13, No. 7, pp. 124-232, plates 4, 5, 7 figs.;
1888, Ann. and Mag. Nat. Hist., ser. 6, vol. i, p. 364; 1888, Proc. Boston Soc. Nat. Hist., vol. 23, p. 391. — HARGITT,
1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 32.

Medusa. — The adult medusa is undetermined, but may possibly be Sarsia flammea. The
most advanced medusae reared by Fewkes had the following characters: Bell rounded,
without apical projection, and in form somewhat fuller than a hemisphere. Size ( ?) Outer
surface of bell besprinkled with nematocysts. Bell-walls of moderate thickness, becoming
thinner near the margin. There are 4 long, equally developed, radially situated tentacles,
with long, tapering basal bulbs which lack ocelli. The 4 radial-canals are broad and straight
and there is a simple, narrow, circular vessel. Manubrium cylindrical and about half as long
as the depth of the bell-cavity. Mouth surrounded by 4 simple lips ( :). The entoderm of
the manubrium is yellow and orange, while that of the tentacle-bulbs is orange. Other parts
of the medusa are colorless.

Fin. 32. — Hydroid of ffyjriehthyi mirus, from Fewkes,
in Bull, \fuseurn Comp. Zool. at Harvard College.

Fish bearing hydroid, and a magnified part of hydroid
itself.

I'oinig medusa. — When first detached from the hydroid the medusa has but two diametri-
cally opposite tentacles, and is more active in its movements than it is when it acquires four
tentacles. This may, however, have been due to the injurious effects of confinement.

Hydroid. — The hydroid was found by Fewkes firmly attached to the side of a small fish,
Seriola zonata. The hydroid colony arises from a basal network of tubes which forms a
plate-like hydrorhiza embedded under the scales of the side of the fish, upon which it is probably
a parasite. This basal network gives rise to tubular gonosomes and also to filiform, flask-
shaped bodies, which recall the spiral zooids of Hydractinia, excepting that they appear to
have a terminal mouth-opening.

The medusa-bearing gonosomes are conical and gradually taper from base to summit,
where there may be a terminal opening. Irregularly distributed side'branches arise from the
main axis of the gonosome, and most of these give rise to medusa-buds. These side branches
are usually simple, although occasionally they branch. They are hollow and in communi-
cation with the entodermal cavity of the main stem.

Clusters of medusa-buds in various stages of development arise from the ends of the
branches. When set free these medusae have but 2 tentacles, but later 2 more tentacles develop.

68

MEDUSA OF THE WORLD.

All parts of the hydroid stock are highly contractile and the gonosomes and zooids are
without chitinous sheaths.

The hydroid is colorless, but the medusa-buds are reddish and orange in color. The
terminal parts of the flask-shaped zooids are also pigmented with reddish-orange.

A single stock of this hydroid was found by Fewkes in Narragansett Bay, Rhode Island,
attached to a fish, Seriola zonata, in August, 1887. The fish was swimming near the surface
and was apparently but little injured by the presence of the parasite.

It is interesting to observe that Alcock, 1892 (Annals and Mag. Nat. Hist., ser. 6, vol. 10,
p. 207, I fig.), has discovered a hydroid of the genus Stylactis which is commensal upon a
fish, Minous inermis. This hydroid, however, produces no medusae.

Genus EUCODONIUM Hartlaub.

Eucodonium, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 71.

The type species is Eucodonium brownei Hartlaub = Dipurena sp. Browne, from the
British Coast.

GENERIC CHARACTERS.

Codonidae with 4 equally developed, radially placed tentacles, each of which terminates
in a distal knob. Stomach mounted upon a gelatinous peduncle. Hydroid unknown.

This genus is distinguished from Sarsia and Slabberia by the gelatinous peduncle upon

which the stomach is placed. It is distinguished from
Dysmorphosa by the terminal knobs of its tentacles and
by its ring-like gonad.

Eucodonium brownei Hartlaub.

Dipurena sp., BROWNE, 1896, Proc. Zool. Soc. London, p. 473, plate 16, fig. 2.
Eucodonium brawnei, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 71, fig. 67.

Bell pyriform, thin-walled, I mm. wide. 4 radially
placed, slender tentacles with small basal bulbs. Ten-
tacles twice as long as the bell-height and with scattered
nematocysts along their lengths, and each with a large,
swollen, nematocyst-beanng knob at outer end of tentacle.
No ocelli. Velum wide. 4 straight, narrow radial-canals.
Stomach short, mounted upon a conical, gelatinous

from Europe, after Hincks in Brit- peduncle. Mouth a simple, round opening at about the

level of the velum. Medusa-buds arise from the sides of

the stomach. The tentacle-bulbs are blackish, stomach and terminal knobs of the ten-
tacles, dull dark-brown. Found at Plymouth, England, in September. Gonads( ?) This
is a young medusa, but we can not refer it to any known mature form.

Genus ECTOPLEURA L. Agassiz, 1862 (sens, amend.).

Tubularia dumortierl, VAN BENEDEN, 1844, Mem. Acad. Belgique, tome 17, p. 50, plate 2.

Eclopleura, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 343. — AGASSIZ, A., 1865, North Amer. Acal., p. 191. — HAZCKEL,
1879, Syst. der Medusen, p. 21. — FEWKES, 1882, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 9, p. 295; 1883, Bull.
Mus. Comp. Zool. at Harvard Coll., vol. n, p. 85, plate I, fig. II. — ALLMAN, 1871, Monog. Tubul. Hydroids, p. 423. —
HINCKS, 1868, British Hydroid Zooph., vol. i, p. 123. — VANHOFFEN, 1891, Zool. Anzeiger, p. 443. — HARTLAUB, 1894,
Wissen. Meeresuntersuch. Komm. Deutsch. Meere Kiel, Abth. Helgoland (l), Bd.l, p. 188; 1907, Nordisches Plankton,
Nr. 12, p. 93. — BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 748.

This genus was founded by L. Agassiz, 1862, the type species being E. dumortieri which
was first described by Van Beneden, 1844, from the coast of Belgium under the name Tubu-
laria dumortien.

GENERIC CHARACTERS.

Codonidae with 2 or 4 simple, unbranched tentacles situated at the bases of 2, or of all,
of the 4 radial-canals. 8 longitudinal rows of nematocysts extend from the 4 tentacle-bulbs
over the surface of the exumbrella to the apex of the bell. The manubnum is short and blunt,
and the mouth is a simple, round opening. The hydroid is Ectopleura, and is closely
related to Tubularia.

FIG. 33. — Eucodonium brownei, after Browne,

in Proc. Zool. Soc. London.
FIG. 34. — Hydroid of Ectopleura dumortien

ANTHOMEDUS.E — ECTO PLEURA. 69

The discovery of Ectopleura pacifica and E. minerva makes it necessary to amend the
generic definition to include medusae with 2 as well as with 4 tentacles.
Ectopleura is distinguished from Zaticlea by its unbranched tentacles.

Ectopleura dumortieri L. Agassiz.

Plate 5, figs. 4, 5; plate 6, figs. I, i', 2.

LITERATURE RELATING TO THE EUROPEAN MEDUSA.

Tubularia dumortieri, VAN BENEDKN, 1844, Mem. Acad. Belgique, tome 17, p. 50, plate I. — JOHNSTON, 1847, Hist. British

Zoophytes, p. 50, plate 7, figs, i, 2.

Ectopleura dumortieri, AC;ASSIZ, L. 1861, Cont. Nat. Hist. U. S., vol. 4, p. 342.
Eclofleura dumortieri, HINCKS, 1868, Hist. British Hydroid Zoophytes, p. 124.
Ectopleura dumortieri, ALLMAN, 1871-72, Monog. Tubularian Hydroid s, p. 424. — BOHM, 1879, Jena. Zcitschrift fiir Naturw.,

Bd. 12, p. 198, taf. 7, figs. 10-13. — HAECKEL, 1879, Syst. der Medusen, p. 22.
Non Ectopleura dumortieri, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 354.
Ectoflfura dumortieri, HARTLAUB, 1894, Wissen. Meeresuntersuch. Komm. Meere Kiel, Helgoland (i), Bd. I, p. 188. — BROWNE,

1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 748.

Ectopkura dumortieri, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 94, figs. 90, 91.
Tubularia dumortieri, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 126 (citation of literature, 1844-48).

LITERATURE RELATING TO THE AMERICAN MEDUSA.

Ectopleura ochracea, A. AGASSIZ, in L. Agassiz's 1862, Cont. Nat. Hist. U. S., vol. 4, p. 343. — AGASSIZ, A., 1865, North Amer.
Acal., p. 191, figs. 320-323. — HAECKEL, 1879, Syst. der Medusen, p. 22. — FEWKES, 1882, Bull. Mus. Comp. Zool. at Har-
vard College., vol. 9, p. 295, plate I, figs. 15, 16, 35, 36. — NUTTING, 1901, Bull.U. S. Fish Commission, vol. 19, p. 373, fig.
82. — HARGITT, 1904, Bulletin Bureau of Fisheries U.S., vol. 24^.32, plate 2, fig. I. — AGASSIZ, A., and WOODWORTH, 1896,
Bull. Museum Comp. Zool. at Harvard College., vol. 30, p. 149, plate 6, fig. 6 (photograph of the medusa).

Ectopleura, sp., HARGITT, 1908, Biol. Bulletin, vol. 14, p. 106, figs. 8-1 1.

The finding of the hydroid of our American Ectopleura "ochracea" leads me to believe
that there are no specific differences between the European and American forms, and that
both should be called Ectopleura dumortieri, this being the older name. Hartlaub, 1907,
however, believes that E. dumortieri is smaller and has somewhat thicker bell-walls than
E. ochracea, but our American medusa varies considerably in these respects, as do also speci-
mens which I captured in the English channel in IQO".

The following description is based upon a study of large numbers of specimens obtained
at Newport, Rhode Island, United States.

Adult medusa. — The bell is pyriform and is about 3 mm. high and 2.5 mm. wide. The
gelatinous substance is very thick and is especially so at the aboral pole. 8 longitudinal lines
of nematocyst-cells extend over the surface of the exumbrella from the basal bulbs of the
4 tentacles to the bell-apex, where they meet in a point. There are 4 equally developed,
short tentacles with large basal bulbs. The outer surface of these tentacles is thickly covered
with nematocyst-cells, and their distal ends are coiled in a close helix. Velum narrow. There
are 4 narrow, straight radial-canals and a simple, narrow, circular tube. The manubrium
is fusiform and extends not more than two-thirds the distance from the inner apex of the
bell-cavity to the level of the velar opening. There is no peduncle, and the mouth is a simple
round opening, the edge of which is armed with clusters of nematocysts. A short, blunt,
axial canal is often seen projecting upward from the stomach into the gelatinous substance
of the apex of the bell. The gonads are developed in the ectoderm encircling the stomach.
In brilliant specimens the tentacle-bulbs are yellow with a red core.

The middle region of the entoderm of the manubrium is lilac-colored, the upper part
delicate yellow, and the lower end red or pink, but very often the medusa is dull purple or
dull brownish-purple.

Hydroid. — The hydroid shown in plate 6, figs. I, I'was found in August, in Newport
Harbor, Rhode Island, at a depth of about 10 fathoms. It is a Tubularian belonging to
the family Hyboconidas Allman, and is apparently identical with 'Tubularia (Ectopleura)
dumortieri of Van Beneden, 1844.

The hydroid is about 25 mm. in length, and was found clinging to the stems of Obelia.

The stems of the Ectopleura are isolated. Most of them are simple and unbranched, but

occasionally they give rise each to a single side branch near the lower end. The lower end of the

Ectopleura is coiled around the stem of the Obelia in a close helix. The stem of the Ectopleura

6

70 MEDUSAE OF THE WORLD.

is covered by a delicate investment of perisarc which displays a number of small annulations
near the upper end. The stem terminates in a single, large polypite, which is broad and
flask-shaped. The mouth is situated at the extremity of a tubular proboscis, and is sur-
rounded by a single verticil of about 16 short, flexible tentacles. In addition to these there is
another verticil of about 24 long, fleshy tentacles at the wide base of the polypite. These long
tentacles are not very contractile. They taper gradually from base to end. The tentacles
at the base of the polypite are about three times as long as are those surrounding the mouth.
The medusa-buds are borne in numbers on short, branching peduncles which arise from the
sides of the polypite in a zone immediately above the basal tentacles. When set free each
medusa has 4 short tentacles, and the 8 longitudinal rows of nematocysts upon the exumbrella
are well developed. The ectoderm of the stem of the hydroid is a delicate, fleshy-yellow and
the entodermal core is flesh-colored pink. The entoderm of the polypite is fleshy-pink and
yellow.

This medusa is very common throughout the summer in Narragansett and Buzzard's
Bays, and on the southern coast of New England east of New Haven, Connecticut. It has
not been taken north of Cape Cod. It has been recorded from Beaufort, North Carolina, and
I found it in Winyah Bay, South Carolina, but it has not been taken either at Charleston,
South Carolina, or at the Tortugas, Florida. It is found in the English Channel and off the
North Sea coasts of England, Scotland, Holland, and Germany. As in America, it appears to
be abundant only in a few localities. I found several specimens of this medusa in the English
Channel in October, 1907, and they appear to be identical in all respects with medusx from
the southern coast of New England, in America. Hargitt has recently discovered the hydroid
at Woods Hole, Massachusetts.

Ectopleura minerva Mayer.
Plate 5, fig. 3.

Ectopleura, sp., FEWKES, 1883, Bull. Mus. Comp. Zool. at Harvard College, vol. II, p. 85, plate I, fig. n.

Ectopleura minerva, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 38, p. 31, plate 16, fig. 38; plate 37, fig. 125.

( ?) Ectopleura dumortieri, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 354.

This medusa has 2 well-developed and 2 rudimentary tentacles, instead of 4 equally
developed tentacles as in the northern species of Ectopleura.

Bell 2.5 mm. high and pear-shaped, with a well-developed apical projection. Bell-walls
of moderate thickness. 8 rows of nematocyst-cells extend from the 4 tentacle-bulbs to the
apex of the bell. There are 2 well-developed tentacles and 2 small tentacle-bulbs. There
are 6 to 9 separate wart-like swellings upon the outer side of each of the 2 large tentacles.
These swellings are crowded with nematocysts. Basal bulbs of tentacles small and without
ocelli. There are 4 straight, narrow radial-canals and a slender, circular vessel. The velum
is well-developed. Manubrium pear-shaped and about two-thirds as long as the depth of the
bell-cavity. It is wider near the middle than at either end. The mouth is a simple, round
opening. A simple, short, conical style-canal extends upward into the apical projection of
the bell.

The entoderm of the manubnum and tentacles is a delicate purple, while the supporting
lamella of the bell is of a decided green. There are a large number of brilliant yellow spots
in the entoderm of the radial-canals and tentacle-bulbs.

This form is rare at Tortugas, Florida. It was found by Fewkes at the Bermudas. A
similar medusa was described by GraefFe, 1884, from Trieste, Mediterranean, under the name
Ectopleura dumortieri.

Ectopleura minerva may prove to be identical with E. pacifica Thornely, from the tropical
Pacific.

Ectopleura pacifica Thornely.

Eclopleura pacifca, THORNELY, 1900, Zool. Results, A. Willey, Part 4, p. 452, plate 44, figs, i, la, Cambridge, England.

Hydroid. — Stems simple, unringed, diminishing in width toward the base, and 20 mm.
high, rising from a creeping stolon which connects the colony. Some of the stems give rise
to stolons near their bases. The polypite is abruptly marked off from the supporting stalk.

ANTHOMEDUS.E — ECTOPLEURA, CORYNITIS. 71

There are two verticils of tentacles of 18 to 20 each. The basal tentacles are very long, and
the oral ones are short, and with knob-shaped ( ?) ends. The medusa-buds are borne in
clusters of from 7 to 9 on branched peduncles, which arise from the body of the polypite
between the two verticils of tentacles. When about to be set free each medusa-bud has 2
well-developed tentacles, thus resembling E. minerva of Tortugas, Florida. E. pacifica is
common in Blanche Bay, New Britain, South Pacific. It is attached to floats, fish-baskets,
etc. Color ( ?) This hydroid may be identical with E. minerva of the tropical Atlantic coast
of North America.

Genus CORYNITIS McCrady, 1857.

Corynitis, McCxADY, 1857, Gymn. Charleston Harbor, Proc. Elliott Soc. Charleston, p. 29. — AGASSIZ, L., i86z, Cont. Nat.

Hist. U. S., vol. 4, p. 340.— AGASSIZ, A., 1865, North Amer. Acal., p. 1 83 .— ALIMAN, 1871, Monog. Tubularian Hydroids,

p. 286.

Coryneles, HAECKEL, 1879, Syst. der Medusen, p. 48.
PlolocniJc, WAGNER, 1885, Wirbelloscn Weissen Meeres, p. 74.
Corynttcs, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 442.

iV<m Corynitis, MURBACH, 1899, Quart. Journ. Mic. Sci., New Series, vol. 42, p. 354 (this is Zanclta}.
(')Modeeria, Protiara (young medusa), HAECKEL, 1879, Syst. der Medusen, p. 47.
(?) Tiaricodon, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.
Non Corynitis, NUTTING, 1901, Bull. U. S. Fish Commission for 1899, p. 372 (this is Zanclta).
Nan Corynitis, HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 42 (this is Zanclea).
Syncorync linvillei (hydroid), HARGITT, 1904, Biological Bulletin, Woods Hole, vol. 7, p. 251, i fig.; Ibid., 1908, vol. 14, p.

101, fig. I.

The type species of this genus is Corynitis agassizii described by McCrady from Charles-
ton Harbor, South Carolina.

GENERIC CHARACTERS.

Codonidae in which the manubnum is cruciform in cross-section. The 4 tentacle-bulbs
have large, abaxial, ectodermal ocelli. The tentacles are unbranched and terminate each in
a knob-like cluster of nematocysts, and the shaft of each tentacle is besprinkled with wart-
like clusters of nematocysts. There are 8 irregular, longitudinal rows of nematocysts upon the
exumbrella, 4 of these being radial and 4 interradial in position. The hydroid is S\ncor\n, .

McCrady, 1857, and L. Agassiz, 1862, described a hydroid which they ascribed to
Corynitis agassizii, but which is certainly very close to, if not identical with, the hydroid
generation of Zanclea. Hargitt, on the other hand, has discovered that Corynitis agassizii
arises from a Syncoryne-\\ke hydroid which he called S. lin-ctllt-i.

In the medusae of Ectopleura, Zanclea, and Corynitis there are longitudinal patches of
nematocysts upon the exumbrella. In Corynitis agassizii the tentacles terminate each in a
knob-like club. In Zanclea dichotoma they terminate in a similar manner, but in addition
they give rise to side branches all arising from the aboral side of the tentacles, and each termi-
nating in a knob-like cluster of nematocysts. In Corynitis, on the other hand, there are
merely scattered wart-like clusters of nematocysts over the shaft of each tentacle. It is possible
that Zanclea arose from some Coryrntis-\]ke medusa; certainly Zancleopsis dichotoma is inter-
mediate in many characters between Corynitis agassizii and Zanclea gemmosa.

However doubtful the relationship may be between Corynitis and the ClaJonemince,
there can be no doubt of its close relationship to Slabberia and Sarsia. The young medusa of
Corynitis can not be distinguished from the young of Slabberia.

Murbach and Nutting are mistaken in their Corynitis medusa, which is clearly Gcmmaria
(Zanclea). The medusa of Corynitis is so clearly described and figured by McCrady as to be
unmistakable. I have frequently taken it in Charleston Harbor, South Carolina, where
McCrady discovered it, and his figure and description enable one instantly to recognize it.
It lacks the feathered tentacles of Gemmarta (Zanclea).

Hargitt, 1904, at first followed Murbach in this erroneous identification of Corynitis,
but his final discovery of the Syncoryni--\\ke hydroid of Corynitis enabled him not only to
correct the mistake, but to establish the close relationship between Corynitis and other Sarsia-
like medusae.

72

MEDUSAE OF THE WORLD.

Corynitis agassizii McCrady.
Plate 5, fig. z.

Corynitis agassizii, McCRADY, 1857, Gymn. Charleston Harbor, p. 30, plate 9, figs. 3-8. — AGASSIZ, L., i86z, Cont. Nat. Hist.

U. S., vol. 4, p. 340. — AGASSIZ, A., 1865, North Amer. Acal., p. 183. — ALLMAN, 1871, Monog. Tubul. Hydroids, p. 287.
Corynetts agassizii, HAECKEL, 1879, Syst. der Medusen, p. 49.

( ?) Corvnetcs agassizii, MAAS, 1893, Ergcb. der Plankton Exped., Bd. 2, K. c., p. 67.

Non Halocharis spiralis (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 239, plate 20, figs. io-ioc.
Coryne agassizii (hydroid), McCRADY, 1857, Gymn. Charleston Harbor, pp. 29, 32.

Non Corvnitis agassizii, HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 42 (this is Gemmaria=ZancIfa).
Non Corynitis agassizii, NUTTING, 1901, Bull. I'. S. Fish Commission for 1899, vol. 19, p. 372, fig. 79 (this is Gcmmaria=

Zanclca*).

Non Corynitis agassizii, MURBACH, 1899, Quart. Journ. Microscop. Sci., New Ser., vol. 42, p. 354 (this is Gemmaria=Zanclea).
Syncoryne linvillei (hydroid), HARGITT, 1904, Biological Bulletin Woods Hole, vol. 7, p. 251, I fig; 1908, Ibid., p. 101, fig. i.

Adult medusa. — Bell miter-shaped and about 2.5 mm. in height. 8 longitudinal rows
of irregularly scattered nematocyst-cells are found upon the sides of the exumbrella immedi-
ately above the circular canal. 4 of these are situated above the tentacles and 4 are inter-
radial in position. The radially situated rows of nematocysts are longer than the interradial.
There are 4 straight, stiff tentacles, which are about three-quarters as long as the bell-diameter.
Each tentacle terminates in a large, knob-shaped swelling, the outer surface of which is thickly
covered with nematocysts. The entodermal core of this swelling is hollow and its lumen

is placed in communication with the gastrovascular system
of the medusa by means of a slender entodermal canal
which extends through the entire length of the tentacle.
Wart-like clusters of nematocysts are scattered over the
shaft of each tentacle. The basal bulbs are large and
hollow, and each one contains a single ectodermal ocellus
upon the outer side. Velum well-developed. There are 4
straight radial-canals and a narrow circular tube. The
large conical apex of the bell is hollow and in direct com-
munication with the stomach of the medusa. The manu-
brium is very large and swollen and fills the greater part of
the bell-cavity. It extends almost to the level of the velar
opening. The mouth is situated at the extremity of a short
oesophagus. It is without prominent lips. The gonads
occupy the greatly swollen sides of the stomach, and their
outer surfaces are smooth. 4 deep, interradial furrows
extend down the sides of the stomach, so that the manu-
bnum is cruciform in cross-section. The entoderm of the
FIG. 35.— Hydroid of Corynitis, after Hargitt, basal bulbs of the tentacles is strawberry-red and that

iqo8, Biological Bulletin. ri 'iiiri I • n

of the terminal knobs of the tentacles is orange-yellow

flecked with red-colored pigment-granules. The manubrium is dull yellow streaked with
strawberry-red and the entoderm of the radial-canals is faint yellow. The ocelli are dark-
brown or orange.

The hydroid is a Syncoryne, and is described by Hargitt. It grows in tufts which are
sparingly branched and 15 to 30 mm. high. The hydranths are vasiform with cone-shaped
proboscis. There are 15 to 30 tentacles distributed over the proximal third of the body of
the hydranth. These tentacles terminate each in a knob-like free end. The pensarc is plain
or with only a slight annulation. The hydrorhiza is reticulated in a loose network. The
medusa-buds are borne upon the body of the hydranth, usually in small clusters at the bases of
the tentacles, each cluster supported upon a single peduncle, the terminal specimen always
maturing first. The medusae, while still attached to the hydroid, have 2 well-developed, dia-
metrically opposed, club-shaped tentacles and 2 small tentacle-buds. There are no ocelli.
There are 4 radial-canals and the umbrella is bell-shaped. In formalin the hydroid is pale-
yellowish, hydranth somewhat brownish, and gonophores reddish-brown or pink. This
hydroid was found by Dr. Henry R. Linville growing upon rocks and piles under "Mill-
dam Bridge "west of Shelter Island, Long Island Sound, New York. It is described by
Hargitt from preserved material under the name Syncoryne linvillei. When set free the
medusae have only 2 tentacles and 2 small, basal bulbs.

ANTHOMEDUSjE — CORYNITIS, SLABBEUIA. 7.1

This medusa is very abundant in Charleston Harbor, South Carolina, during the summer
and early autumn. It is also common in Great Peconic Bay, Long Island Sound, and at
Woods Hole, in September. It has not been taken north of Cape Cod, Massachusetts, or
at the Tortugas, Florida.

Corynitis arcuata Haeckel.
Coryneies arcuala, HAECKEL, 1879, Syst. der Medusen, p. 49.

This variety, or species, is distinguished from Corynitis agassizii by its long, narrow
bell, about twice as high as it is broad, and by its long, thin tentacles. The nematocysts upon
the tentacles are smaller and more numerous than they are upon the tentacles of C. agassizii.
The stomach is small and bell-shaped, with a short oesophagus which does not extend beyond
the velar opening. Haeckel gives the bell-height as 6 and the breadth 10 mm.; but judging
from his description of the medusa these figures must be erroneously stated and should be
reversed.

It is found off the coast of Brazil. Haeckel describes it from a single preserved specimen.

Corynitis (?) coerulea.
Tiaricodon cxruleus, BROWNE, 1902, Annals Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

This medusa is very briefly described by Browne, without figures. The gonads are not
radially separated and there is nothing in Browne's description to indicate that the medusa
belongs to the Tiarinae. Its characters appear to be those of Corynitis.

Umbrella bell-shaped, 25 mm. high, 24 mm. wide, with a rounded summit. The exum-
brella is smooth without nematocysts. 4 fairly stout, radially situated tentacles tapering to a
point. Their basal bulbs are large and cylindrical, a little longer than broad, and with an
ocellus. Velum narrow. The manubrium is placed upon a short, broad peduncle. It is quad-
rangular with 4 perradial folded lips about as wide as the stomach. The mouth extends
nearly to the level of the velar opening. The gonads surround the stomach and extend outward
over the peduncle as 4 perradial lobes. Found at Stanley Harbor, Falkland Islands. The
young medusa resembles a Sarsia. The color is not stated by Browne.

Genus SLABBERIA Forbes, 1846.

Slabbcria, FORBES, 1846, Ann. and Mag. Nat. Hist., vol. 18, p. 286; 1848, British Naked-eyed Medusx, p. 53. — FORBES and

GOODSIR, 1853, Tran. Roy. Soc. Edinburgh, vol. 20, p. 311.
Dipurrna, McCRADv, 1857, Proc. Elliott Society, Charleston, p. 135. — AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341. —

AGASSIZ, A., 1865, North Amer. Acal., p. 180.

Djpurtna + Bathycodon, HAECKEL, 1879, Syst. der Medusen, pp. 23, 26.
Dipurfna, BROWNE, 1898, Proc. Zool. Soc. London for 1897, p. 816.

Dipurena+Dipurella, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, pp. 30, 31.
Dipurtna, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 442. — BROWNE, 1905, Report on Pearl Oyster Fisheries, Gulf of

Manaar, Supplementary Report 27, Roy. Soc. London, p. 133.
S/abbcria (in part), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.
Slabberia (in part), HAECKEL, 1879, Syst. der Medusen, p. 23.
Purena+ S/abbcria, H\RTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 55, 62.

In 1846 Forbes described a medusa under the name Slabl>,-ri<i hultfrata. It had 4
radial-canals with a swollen region upon each canal. In other respects it resembled Sarsiti,
excepting that each of the 4 tentacles terminated in a knob-like extremity. Forbes believed the
swelling on the radial-canals to be gonads and established the genus Slabbtria with the
following designation: "Umbrella campanulate; ovaries four, linear, in the course of the
four simple, gastrovascular canals; peduncle proboscidiform, highly extensile, oral orifice cir-
cular; a marginal tentacle springing from an ocellated bulb and terminating in a colored
globular body, placed opposite each of the gastrovascular canals."

Forbes laid especial stress upon the "gonads" upon the radial-canals, and he believed the
manubrium to be simple and tubular, and failed to find collar-like swellings upon it. Haeckel,
1879, p. 241, and Browne, i8g8, p. 816, find, however, that the gonads in Forbes' s medusa are
developed in separated ring-like annuli upon the manubrium and Browne, who sectioned the
supposed "gonads," finds that they contain entodermal cells crowded with small nuclei, and
they are doubtless mere glands similar to those seen upon the radial-canals of many Antho-

74

MEDUSA OF THE WORLD.

Tabular Description of the Medusa of Slabberia.

Slabberia hal-

S.strangulata =

S. strangulata

S.catenata Forbes,

S. ophiogaster=

S . pyramis= Bathy-

terata Forbes,

Dipurena

var. fragilis

1853= Dipurena

Dipurena

codon pyramis

I846.1

strangulata

Mayer.3

dolichogasta

ophiogasler

Haeckel, 1879.°

McCrady,

Haeckel, 1879,

Haeckel, 1879.

i857.2

= D. picta Mayer,

= Sarsia stran-

1900= Sarsia

gulata Allman,

clavata Kefer-

1 87 1 .5

stein, i862.4

Shape and

Ellipsoidal.

Ellipsoidal to

Ellipsoidal.

Ellipsoidal.

Globular.

Pyramidal.

size of bell

8 high, 6 wide.

hemispherical.

4 high, 3.5

24 high, i. 6 wide.

5 high, 4 wide.

4-sided. 5 high,

in mm.

2 to 4 high,

wide.

5 wide. 4Hnes of

4103.3 wide.

nettle-cells ex-

tended up ex-

umbrella in 4

principal radii.

Character of

Large with an

As in S.halter-

Very small.

As in S. halterata.

Poorly developed,

Ocelli ?

the 4 ten-

abaxial ocellus.

ata.

with abaxial

tacle-bulbs

ocelli.

Length of

2 r

1.5 r, quite

2 r very slender.

2r±

2r +

4r +

each ten-

thick.

tacle in

terms of

bell-ra-

dius (r).

Number and

i large, terminal,

Single knob-like

As in S. strangu-

Proximal halves of

Tapering. Outer

Each tentacle ter-

position of

club-shaped

end and with

lata.

tentacles smooth.

parts thickly

minates in

nettle-

knob, i to 2

nettle-warts

Outer halves

covered with

"suctorial cup."

warts on

rings at base

on shaft of

ringed with 15 to

prominent par-

each ten-

of knob.

each tentacle.

3onematocyst

tial rings and

tacle.

swellings.

spirals contain-

ing nettle-cells.

Character

Narrow, straight,

Narrow, straight,

As in S. strangu-

As in S. strangulata.

As in S. strangu-

Edges of radial-

of the 4

each with swol-

without "glan-

lata.

lata.

canals lined

radial-

len(glandular?)i dular" swell-

with gland-cells.

canals.

enlargement at

ings.

two-thirds

length of canal

above margin.

Length of

4 to 71-.

4 to 7 r.

2 r

4 to 7 r.

8 to 12 T.

4r+

manubrium

in terms of

bell-radius

Number of

2 or 3. I or 2

2. Small ring

i. As in S.

2 or 3 as in S. hal-

4 to 6 short,

8 separated rings-

ring-like

small genital

near middle,

strangulata.

terata. Preceding

swollen, sepa-

gonads

rings below

and longer

development of

rated by short

which en-

middle, and a

outer ring.

gonads medusa-

intervals.

circle

longer one

buds are borne on

manu-

near outer end

manubrium (see

brium.

of manubrium.

text).

Color.

Entoderm of ten-

Variable shades

Dull-yellow en-

Stomach yellow or

Asin S.catenata.

Stomach canals

tacle-bulbs

of yellow, red,

toderm of

red. Gonads and

Ocelli black.

and tentacles

and tentacle-

and green.

stomach and

tentacles light-red.

light-green.

knobs red-

Ocelli black.

tentacles.

Mouth, tentacle-

Mouth, tenta-

dish-brown;

Ocelli black.

bulbs, and apex of

cle-bulbs and

of manubrium

stomach bright-

terminal "an-

yellowish-

yellow, red, and

chors" darker

brown.

purple. Ocelli

green.

Ocelli black.

deep-purple brown.

Where found.

Southern coasts

Coast of United

Tortugas,

West coast of Scot-

Southern coasts of

Mediterranean,

of England

States from

Maldive Is-

land to Mediter-

England and

Corfu; from

and Ireland.

South Caro-

lands, Indian

ranean.

Ireland to Med-

depth of 20

Rare.

lina to Cape

Ocean, Bige-

iterranean. ( ?)

fathoms, April,

Cod.

low.

Cevlon (Browne).

1877, Haeckel.

'Distinguished by 4 small, isolated (glandular?) bodies upon its 4 radial-canals. Hartlaub, 1907, believes these to be gonads
which become mature only in old medusae.

2 Distinguished by its quite thick tentacles, each ending in a single, knob-like cluster of nematocysts.

3 Separated from S. strangulata only by its long, slender tentacles and uniform dull-yellow color.

4Hydroid a Syncoryne. Medusa distinguished by numerous total or partial rings of nettling cells which are developed over

outer halves of tentacles. When young medusa-buds are borne on manubrium of free medusa.
5Is this a fully developed, mature condition of 5. catenata(t).
6 Distinguished by terminal "suctorial cups" on its tentacles and numerous glands (?) which line radial-canals.

ANTHOMEDUS^E — SLABBEIIIA. 75

medusae. On the other hand, Browne found the gonads upon the manubrium, arranged in
2 or more isolated ring-like swellings. In 1853 Forbes and Goodsir describe another species
of Slabberia, S. catenata, which lacks glands upon its radial-canals, although Forbes states that
they exist but are "scarcely to be traced."

In 1857 McCrady described a medusa under the name Dipurena strangulata, and in this
the 4 radial-canals are simple, but the manubrium is tubular and encircled by 2 separated,
ring-like gonads.

It is evident that Slabberia of Forbes is similar in all respects to Dipurena, excepting for
the problematical swellings on the radial-canals. It seems best to drop Dipurena and retain
the older name Slabberia. Swellings similar to those on the radial-canals of S. halterata are
seen on the radial-canals ot other Anthomedusae, such as Lymnorea alexandri and Dysmor-
phosa dubta. In L. alexandri the swellings are glandular and they are probably of similar
function in the Slabberia ot Forbes. Hartlaub (1907), however, states that these swellings are
gonads which develop only in old medusae. He does not state, however, that he sectioned the
medusa, and the matter appears to me to be still in doubt.

GENERIC CHARACTERS.

Codonidae in which the manubrium is tubular and is encircled by two or more separated
collar-like gonads. In other respects the medusa resembles Sarsia. The 4 radial-canals are
simple and without gonads. The hydroid is Syncoryne.

In all known species of Slabberia each of the 4 tentacles terminates in one or more knob-
like clusters of nematocysts.

Haeckel, 1879, p. 26, instituted the genus Bathycodon for a medusa resembling Dipurena,
but with short, blindly-ending diverticula upon its radial-canals. We define Slabberia as
equivalent to Dipurena+Bathycodon as defined in Haeckel's System.

Haeckel suppresses the generic name Slabberia on the ground that it was incorrectly
defined by Forbes, but if we are to retain only the correctly defined genera we must at once
drop nearly all of those defined previous to 1850, and even the majority of modern generic
names must vanish! As knowledge increases it appears to me far less confusing to amend and
retain old names rather than to drop them and substitute unfamiliar designations for familiar
objects.

Hartlaub, 1907, would distinguish Purena, with nettle-warts (not rings) upon the tentacles,
whereas Slabberia he would confine to include medusae having tentacles ringed with nema-
tocysts. The distinction between broken rings ("warts") and complete rings appears to me to
be too slight to be of generic value, but is an important specific distinction.

Slabberia halterata Forbes.

Slabberia halterata, FORBES, 1846, Annals, and Mag. Nat. Hist., vol. 18, p. 286; 1848, British Naked-eyed Medusae, p. 53, plate

6, fig. i. — AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.
Dipurena halterata, HAECKEL, 1879, Syst. der Medusen, p. 24.
Dipurena halterata, BROWNE, 1896, Proc. Zool. Soc. London, p. 473; 1897, Ibid., p. 816; 1898, plate 49, figs. 2-iA. — BEDOT,

1905, Revue Suisse de Zool., tome 13, p. 133 (citation of literature 1846-49).
Slabberia halterata, HART-HUB, 1907, Nordisches Plankton, Nr. 12, p. 64, figs. 61, 62.

Bell ellipsoidal, 8 mm. high, 6 mm. wide, with thick walls. Bell cavity about three-fourths
as deep as the height of the bell. 4 slender tentacles with well-developed basal bulbs, each with
an abaxial, ectodermal ocellus. Each tentacle terminates in a large, elongate, swollen tip with
one or two complete rings of nematocysts above it. 4 radial-canals straight and narrow, and
each exhibits I or 2 small (glandular?) swellings (gonads?) at about one-third the distance
between the inner apex of the bell-cavity and the margin. Hartlaub, 1907, believes these
swellings to be true gonads which become mature only in old medusae.

The stomach-cavity extends upward a short distance above the base of the manubrium
into the gelatinous substance of the apex of the bell.

The manubrium is 4 to 7 times as long as the radius of the bell. It is slender and tubular
and there is a small, spindle-shaped stomach near its distal end. The mouth is at the end of a
short, conical neck and is a simple, round opening.

76 MEDUSAE OF THE WORLD.

2 or 3 ring-like gonads girdle the manubrium. The two upper gonads are short, the
uppermost one being near the middle of the manubrium, and the next lower separated from it
by a short interval. The lowermost (distal) gonad is about twice as long as either of the others,
and it is adjacent to and above the stomach.

The entoderm of the basal bulbs and terminal clubs of the tentacles is reddish-brown and
of the manubrium yellowish-brown. The ocelli are black.

This medusa has been taken at infrequent intervals off the southern coasts of England and
Ireland. The best modern descriptions are given by Browne, 1898, and by Hartlaub, 1907.

It is distinguished by the 4 small swellings on the 4 radial-canals. According to Browne
these are probably glandular, but Hartlaub regards them as gonads which develop later than
the gonads upon the manubrium. Hartlaub, apparently, has not sectioned the medusa, while
Browne bases his conclusions upon a study of sections.

Slabberia strangulata Haeckel.
Plate 7, figs. I, 2, and 3.

Dipurtna strangulata, McCRADY, 1857, Gymn. Charleston Harbor, p. 33, plate 9, figs. I, 2.

Dipurena cervicata, McCRADY, 1857, Gymn. Charleston Harbor, p. 34.

Dipurena conica, A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Dipurcna cervicata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Difurena strangulata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Difurena conica, AGASSIZ, A., 1865, North Amer. Acal., p. 181, figs. 301, 305.

Dipurena strangulata= Slabberia strangulata, HAECKEL, 1879, Syst. der Medusen, p. 23.

Dipurena conica, HAECKEL, 1879, Syst. der Medusen, p. 24.

Dipurena strangulata, FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 155, plate 4, fig. 5. — BROOKS, 1882, Studies Johns

Hopkins Univ. Biol. Lab., vol. 2, p. 136.

Dipurena conica, VERRILL, 1873, Report Commiss. Fish and Fisheries for 1871-72, p. 735.
Dipurella clavata (immature), HARGITT, 1904, Bulletin of the Bureau of Fisheries, vol. 24, p. 31, plate I, fig. 3.
Dipurena conica, NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 373. — HARGITT, 1901, American Naturalist,

vol. 35, p. 578, fig. 34.
Purena strangulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 55, fign. 51-53.

In 1857 McCrady described those which he believed to be two distinct species of Difurena,
D. strangulata and D. cervicata, from Charleston Harbor, South Carolina. In 1862-65,
A. Agassiz described that which he believed to be another species, from Buzzards' Bay, under
the name of D. conica. A careful study of Dipurena in Charleston Harbor, in September, 1897,
when the medusae were exceedingly abundant, has convinced me that all of these forms are one
and the same species. McCrady found only two individuals of D. strangulata and one of
D. cervicata, and thus A. Agassiz was misled into the assumption that the northern form was a
distinct species. The shape of the bell is, however, very inconstant, and the length of the
manubrium in the Charleston medusae is subject to a wide range of variability.

Adult medusa (plate 7, fig. i). — The bell is quite variable in shape, for in some individ-
uals it is hemispherical, while in others it is ellipsoidal and higher than a hemisphere. All
gradations between these two extremes are observed. In a large individual in which the bell
was hemispherical it was 4 mm. in diameter; while another having an ellipsoidal bell was 4 mm.
in height and 3 mm. in width. There are 4 stiff tentacles, one at the base of each radial-
canal. Each tentacle terminates in a single, knob-like swelling which is covered with nema-
tocysts. The entodermal core of this swelling is hollow, and its lumen is placed in communi-
cation with the general gastrovascular system of the medusa by means of a narrow thread-like
canal which extends through the entire length of the tentacle. The basal bulb of each tentacle
is large and there is a single, ectodermal ocellus mounted upon a short pedicel, upon the outer
side of the bulb. The tentacles are all of equal length, and each one is about three-fourths as
long as the bell-diameter. The velum is wide. There are 4 straight, simple radial-canals
and a narrow, circular tube. The gastric cavity projects upward a short distance into the
gelatinous substance of the bell at the point of intersection of the 4 radial-canals. The manu-
brium is always long and increases in length with advancing age. In mature medusae it projects
far beyond the velar opening and varies in length from 5 to 10 mm. It is tubular and highly
contractile, and the mouth is a simple, round opening. Four distinct regions may be distin-
guished upon the manubrium of the mature medusa. First, a long, narrow, tubular region,
extending from the inner apex of the bell-cavity to about the level of the velar opening, then a
short, fusiform, swollen region, then a short, narrow, tubular region similar in size to the

PLATE 7.

Fig. I. Slabberia strangulata, mature male. Charleston Harbor, South

Carolina, September 8, 1897.
Fig. 2. Slabberia strangulata, half-grown medusa. Charleston Harbor,

South Carolina, September 1 1, 1897.
Fig. 3. Slabberia strangulata, young medusa. Charleston Harbor, South

Carolina, September 7, 1897.

Fig. 4. Slabberia strangulata var. fragilts. Tortugas, Florida.
Fig. 5. Zanclea gemmosa, young medusa. Agassiz Laboratory, Newport,

Rhode Island, September 17, 1892.

Drawn from life, by the author.

ANTHOMEDUS^E — SLABBERIA. 77

proximal part of the manubrium, and finally at the distal end a long, club-shaped, cylindrical
region, at the extremity of which the mouth is situated. The 2 genital organs are found
within the 2 swollen regions of the manubrium. The color of these medusae is quite variable,
the entoderm of the manubrium and tentacles being either yellow, or yellow and red, or red,
or green and red. In the young medusas the entoderm of the manubrium is usually yellow, and
of the tentacles yellow dotted with red granulations; while in the mature medusa the color
of the entoderm is usually green or yellowish-green.

Young medusa (plate 7, fig. 3). — In the youngest medusa observed, the bell is cylindrical
with vertical sides and a slight apical projection. The gelatinous substance is thin. There are
2 diametrically opposed tentacles each of which terminates in a single, large, nematocyst-
covered knob. The other 2 tentacles are as yet undeveloped and are represented by mere
basal bulbs. There are 4 ocelli, one upon each basal bulb. The manubrium is a simple, short,
conical tube and the mouth is a simple, round opening.

When the medusa is I mm. in height, a constriction appears near the proximal end of the
manubrium, which thus becomes divided into an upper and lower swollen region. As develop-
ment proceeds this constriction becomes more marked and finally the manubrium assumes
the adult shape; the upper and lower swellings of the young proboscis corresponding to the two
swollen regions of the manubrium of the adult medusa. Hargitt, 1904, describes the young
medusa under the name of "Dtpurella clavata."

This medusa is exceedingly abundant in August and September in Charleston Harbor,
South Carolina. It has been found by Brooks at Beaufort, North Carolina; and by A.
Agassiz and Verrill in Buzzards' Bay, Massachusetts. It has not been taken north of Cape
Cod, Massachusetts, and has not been observed at the Tortugas, Florida.

Slabberia strangulata var. fragilis.
Plate 7, fig- 4-

Difurena fragilis, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 29, plate 17, fig. 41.
(?) Difurena fragilis, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251.

Bell 4 mm. high and halt-egg-shaped, with moderately thick walls, becoming thinner
near the margin. There are 4 long, slender tentacles, each about as long as the bell-height.
These terminate each in a single knob armed with nematocysts. The tentacle-bulbs upon
the bell-margin are not large, but each one bears a single, black ocellus in the ectoderm of its
outer side. Velum wide. There are 4 straight, narrow radial-canals and a slender, circular
vessel. Manubrium about twice as long as bell is high. It is slender and tubular, with two
separate, swollen regions containing the gonads. The entoderm of the manubrium and of
the basal bulbs of the tentacles is dull yellow, while the entoderm of the terminal knobs of
the tentacle is a faint orange.

This variety is found at Tortugas, Florida, in June. Bigelow, 1904, found a colorless
Dipurcna at Suvadiva Atoll, Maldive Islands, Indian Ocean, which he believes is probably
identical with Dipurena fragilis.

S. fragilis may be distinguished from other American Dipurenae by its \ong,very slender
tentacles, and the dull-yellow color of its entoderm.

Slabberia catenata Forbes and Goodsir.

Plate 8, figs. 8 and 9.

Slabberia catenata, FORBES and GOODSIR, 1853, Trans. Roy. Soc. Edinburgh, vol. 20, p. 311, plate 10, figs. ya-^e.

Difurena dolichogaster, HAECKEL, 1864, Jena. Zeltschrift fur Naturw., Bd. i, p. 337; 1879, Syst. der Medusen, p. 25, taf. 2,

figs. 1-7.

Difurena calenata, HAECKEL, 1880, Syst. der Medusen, p. 655.

Difurena fertilis, METSCHNIKOFF, E. and L., 1870, Verhandl. Gesell. Freunde Nat. Moskau, tome 8, p. 343, taf. 3, figs. 1-5.
Sarsia dolichogaster, SPAGNOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 18, tav. 2, fig. 3 (from Naples, Italy).
Difurena dolichogaster, CHUN, 1895, Bibliotheca Zoologica, Heft 19, p. 6, taf. i, figs. 1-4. *
Difurena catenata, MONTICELLI, 1897, Atti Accad. Sci. Torino, vol. 32, p. 888, I fig. (abnormal specimen from the Gulf of

Cagliari, with a manubrium which gives off 2 side branches, each with mouth and gonad).
( ?) Sarsia clavata, KEFERSTEIN, 1862, Zeitschrift fur wissen. Zool., Bd. 12, p. 27, taf. 2, figs, i, 2.
( ?) Sarsia, sp., ALLMAN, 1871, Monog. Tubul. Hydroids, p. 83, fig. 37.
(?) Sarsia clavata, HAECKEL, 1879, Syst. der Medusen, p. 19.

( ?) Sarsia clavata (Syncoryne clavata), GRAEFFE, 1884, Arbeit. Zool. Inst. \Vien, Bd. 5, p. 351 (first description of the hydroid).
Difurena ficta, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 29, plate 18, figs. 45, 46.
(? Sarsia clavala)^ Slabberia catenata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 51, 63, figs. 46, 59.

78 MEDUSAE OF THE WORLD.

DESCRIPTION OF AMERICAN SPECIMENS.

Bell high with straight, vertical sides and flatly-rounded top, about 3 mm. high and 1.5
mm. wide. Bell-walls very thick and of a rigid, gelatinous consistency. Bell-cavity about
two-thirds as deep as the total height of the bell itself. There are 4 radially situated, slender
tentacles, all of the same length and not quite as long as the bell-height. Each ot these ten-
tacles terminates in 3 to 5 bulb-shaped, ring-like swellings containing nematocysts. The
basal bulbs of the tentacles are large and swollen, and each of them bears a dark-purple,
ectodermal ocellus upon its outer side. There are 4 straight, narrow radial-canals and a
simple, slender, circular vessel. The velum is quite narrow. The manubrium is a slender
tube about 5 mm. long; and it exhibits two separate, annular, swollen regions marking
the places where the gonads are situated. The mouth is a simple, round opening. The ento-
derm of the manubrium and of the tentacle-bulbs is of a beautiful custard-yellow, while the
entoderm of the terminal series of knobs on the tentacles is of a rich port-wine color.

Several specimens were found in a surface-tow at Tortugas, Florida, early in August,
1898, and in July, 1906.

DESCRIPTION OF EUROPEAN SPECIMENS.

For details of the general characters of the European medusa, see tabular description
of the medusae of Slabberia.

Forbes and Goodsir, 1853, describe this medusa from the western coast of Scotland, at
Tobermory and Loch Laigh.

Keferstein, 1862, and Allman, 1871-72, describe that which I doubtfully believe to be
this medusa from the coasts of France and England. It is the immature condition before
the appearance of gonads, while medusa-buds are still being produced upon its manubrium,
and before the development of nettle-rings upon its tentacles. Chun, 1895, has made a most
careful study of this budding process in specimens from the Mediterranean.

Haeckel, 1879, describes this medusa from the Mediterranean, but did not observe it
in the stage wherein medusa-buds are being produced. An identical medusa is found at
Tortugas, Florida.

Chun, 1895, has made a careful study of the budding process. When the medusa^is
young the manubrium is slender and cylindrical and without gonads. Medusa-buds develop
at this stage. These budding medusae develop in a helical line at regular intervals down
the sides of the manubrium. The oldest of the primary series of medusa-buds is nearest the
base and the youngest nearest the mouth of the manubrium.

These primary medusa-buds are attached by narrow pedicels to the sides of the manu-
brium, and a second series of medusa-buds develop upon the sides of these pedicels, before
the primary medusae are set free. After the primary medusae are set free the secondary medusa-
buds complete their development. Thus the manubrium may present various appearances;
the oldest medusa-buds at any one time may be above, or below, or anywhere upon the manu-
brium. Before they are set free the budding medusas begin to develop a secondary genera-
tion of medusa-buds upon their manubria.

The hydroid of this medusa is described by Graeffe from Trieste, Adriatic Sea. GraefFe
succeeded in rearing the hydroid from the eggs of the medusa. It is a Syncoryne. The poly-
pites are cup-shaped, only 2 to 3 mm. long, and have 4 circlets of 4 tentacles each. The
tentacles are short and stiff, and each one ends in a nematocyst-knob. The short stems of
the hydroid are invested by an irregularly ringed perisarc. The hydrorhiza forms a mass of
profusely branched anastomosing stolons. The medusa-buds arise from the sides of the poly-
pites between the 2 lowest circlets of tentacles. When set free the medusa has a short man-
ubrium and 4 tentacles. The tentacles have terminal nematocyst-knobs and a nettling ring.

The eggs are laid in May and June and the mature hydroid is found in October at Trieste.

Chun is of the opinion that Sarsia clavata is identical with "D. dolic hogaster" ; but
Hartlaub regards S. clavata as being distinct and a true Sarsia. The absence of a series of
rings upon the tentacles of S. clavata inclines me to agree with Hartlaub that it is a Sarsia
and not to be confused with Slabberia catenata. Further studies appear to be required before
any final opinion can be formed upon this case.

ANTHOMEDUS.E — SLABBERIA.

79

Slabberia ophiogaster.

Sarsia strangulate, ALLMAN, 1871-72, Monograph Tubularian Hydroids, p. 46, fig. 17.

Dipurena ophiogaster=Tetrapurena ophiogaster, HAUCKEL, 1879, Syst. der Medusen, p. 25.

Dipurena ophiogaster ( ?), BROWNE, 1905, Pearl Oyster Fisheries, Gulf of Manaar, Roy. Soc. London, Suppl. Report 27, p. 133,

plate 2, figs, i, 2 (from Galle Bay, Ceylon).
Purena strangulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 55, figs. 51-53.

This medusa is found off the southern coasts of England and Ireland, and an apparently
identical species is described by Browne from Ceylon. Dr. Lobianco kindly permitted me
to study a specimen, which I consider to be of this medusa, from Naples, Italy.

For details, see tabular description of the medusae of Slabberia.

FIG. 36. — Slabberia ophiogaster. Drawn by the author, from a specimen collected by Dr. Lobianco in the Bay of Naples.
FIG. 37. — "Purena strangulata" (5. ophiogaster), after Hartlaub, in Nordisches Plankton.

In the specimen in Dr. Lobianco's collection from Naples, the bell is 5 mm. high, 4 mm.
wide; oval, with thick walls. The manubrium is 13.5 mm. long and has 2 gonads, one in the
middle of its length and one adjacent to the stomach. The 4 tentacles are each 2.5 mm. long,
and their outer halves taper to a point and are covered with broken, partial rings of nettle-
cells. The proximal halves of the tentacles are smooth and cylindrical. The tentacle-bulbs
are swollen below the bell-margin and have ocelli.

Slabberia pyramis.
Baltivcodon pyramis, HAECKEL, 1879, Syst. der Medusen, p. 26.

This Mediterranean medusa is distinguished from other species of Slabberia by the
terminal "suction cups" upon its tentacles and by the numerous "glandular" organs which
line the sides of the radial-canals. For details see the tabular description of medusae of
Slabberia.

MJ MEDUSAE OF THE WORLD.

Genus MARGELOPSIS Hartlaub.

Nemopsis (hydroid only), MCCRADY, 1857, Gymn. Charleston Harbor, Proc. Elliott Society of Charleston, vol. I, p. 163, plate

10, figs. 4-7.
Marsehftshi HARTLATB, 1897, Hydromedusen Helgolands, Wissen. Meeresuntersuch., Biolog. Anstalt auf Helgoland, Neue

Folge, Bd. 2, p. 48z; Ibid., 1904, Bd. 5, p. 99; 1899, Nachrich. k. Gesell. der Wissenschaften Gottingen, math.-phys.

Klasse, p. 219; Zool. Centralbl., Bd. IO, p. 28; 1907, Nordisches Plankton, Nr. 12, p. 88. — BROWNE, 1903, Bergens

Museums Aarbog, No. 4, p. 10. — MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, pp. 43, 73.

GENERIC CHARACTERS.

Anthomedusae with simple, round mouth-opening without oral tentacles, or cruciform
lips. With 4 radially placed clusters of marginal tentacles and 4 radial-canals. The ring-
like gonad encircles the stomach. The hydroid is the pelagic Margelopsis.

The tentacles of the hydroid are confined to definite whorls. In 1857, McCrady described
a free-floating hydroid from Charleston Harbor which has two whorls of tentacles, and between
them a number of medusa-buds arise from the sides of the hydranth. When set free the
medusas bore some resemblance to Nemopsis, and McCrady believed them to be the young
of Nemopsis gibbesn = Nemopsis bachei L. Agassiz. We now know this to have been a
mistake, for Hartlaub, 1899, discovered that actinula larvae develop upon the sides of the
manubrium in Margelopsis haeckehi. Then these actmulae are set free with two whorls of
tentacles, and medusa-buds develop from the sides of the hydranth between the whorls, and
are set free, as in McCrady's hydroid, which is, indeed, a species of Margelopsis.

In 1904 Hartlaub found another free-floating hydranth (Margelopsis stylostoma) which
is apparently detached from Tiarella singularis F. E. Schulze. It is apparent that the Siphon-
ophoras may have been derived from hydroid forms having a similar life-history.

The medusas of this most interesting genus appear to form a connecting link between
the Codonidae and Margelidae of Haeckel. They resemble the Codonidae in their simple
mouth-opening without oral tentacles or prominent lips, and in the single ring-like gonad
surrounding the manubrium. On the other hand, they resemble the Margelidae in having
the marginal tentacles grouped into 4 radially situated clusters. There are, however, no
tentacular ocelli such as are seen in the Margelidas.

The Atlantic species are all very closely related and may prove to be but one. The
chief distinctions are the number of tentacles upon each tentacle-bulb. For example: M. hart-
lanlm Browne has 2; M. haeckeln Hartlaub has 3 to 4; M. gibbesi Hartlaub has 510 6.

The medusa of M. stylostoma, and the hydroid of M. hartlaubii are unknown, and it is
possible that these two may be phases in the life-history of one and the same animal.

The so-called "Perigonimus sulphureus" Chun, 1889 (Sitzungsber. Akad. Wissen. Berlin,
Jahrg. 1889, p. 524), and Steche, 1906 (Zool. Anzeiger, Bd. 31, p. 30, 2 figs.), is probably
not a Perigonimus, but may give off pelagic hydranths. Chun found this hydroid grow-
ing upon a living shell of ffyaliea in the Canary Islands. One portion of the net-like
hydrorhiza gives rise to isolated polypites, and another portion to the medusa-buds. The
hydranths arise directly from the hydrorhiza and are unbranched. They are barrel-shaped,
large, and with an oral circlet of 6 to 9 short, blunt tentacles. One of the hydranths was sharply
constricted by a ring-furrow in the middle of its length as if the oral end were about to be set
free and to become pelagic (Steche). The medusa-buds also arise singly from the hydrorhiza,
each of the medusa-buds being placed upon a short, unbranched peduncle. When set free
the medusas have 4 short tentacles with swollen basal bulbs, and the manubrium does not
project beyond the velar opening. The entoderm of the hydroid and medusa is sulphur-
yellow. The periderm of the hydroid is smooth and has none of the foreign bodies attached
to it such as are seen in Perigonimus. According to Steche the colony feeds upon the eggs
which are set free by the Hyaliea on which it grows.

Margelopsis haeckelii Hartlaub.

Margelopsis hxckelii, HARTLAUB, 1897, Wissen. Meeresuntersuch. Kommis., Meere Kiel, Helgoland, Neue Folge, Bd. 2, p. 482,
taf. :6t, fign. 12-18; 1899, Nachricht. kg!. Gesell. Wissen. math.-phys. Klasse, Gottingen, Jahre 1899, p. 219, 3 fign.;

1907, Nordisches Plankton, Nr. 12, pp. 89, 91, fign. 84-86 (best published description of hydroid and medusa). — MULLER,

1908, Zeit. fur wissen. Zool., Bd. 89, pp. 43, 73, taf. 5, fign. 18, 19.

PLATE 8.

Fig. I. Zancleopsls dichotoma. Tortugas, Florida, May 28, 1899.

Figs. 2 and 3. Gemmaria costata ( ?), hydroid and medusa. Tortugas,
Florida, June, 1899. This is probably the hydroid of Zanclea
costata.

Figs. 4 and 5. Zanclea gemmosa, young medusa. Figure 5 shows details of
nematocyst-capsules of the tentacles. Agassiz Laboratory, New-
port, Rhode Island, June 19, 1893.

Fig. 6. Zanclea costata. Tortugas, Florida, May 15, 1906.

Fig. 7. Zanclea costata. Details of nematocyst-knobs of tentacles.

Figs. 8 and 9. Slabberia catenata. Tortugas, Florida, August, 1898.
Figure 9 shows end of one of the tentacles of the medusa shown in
figure 8.

Drawn from life, by the author.

ANTHOMEDUS.E — MARGELOI'SIS.

SI

Medusa. — Bell pyriform with a flat apex 1.3 times as high as wide. 2 mm. high. Gelat-
inous walls quite thick. 3 or 4 tentacles upon each of the 4 marginal tentacle-bulbs. These
tentacles are ringed with nematocysts. No ocelli. 4 narrow, straight radial-canals and a
wide axial-canal above the stomach. Manubnum wide, spindle-shaped, two-thirds as lout;
as the depth of the bell-cavity. The upper third ot the manubnum has no gonad, but the
lower part is incased in a tubular genital organ which bears planulae. Tentacle-bulbs
brown. Manubrium with dark-brown pigment granules. Found at Helgoland, German
Ocean, in July and early in August.

Hydroid. — The planulae which develop upon the manubrium of the medusa grow into
actinula larvae before being set free. The oral pole of the actinula is blunt and is adjacent
to the wall of the stomach of the medusa, while the aboral pole projects outward from the
stomach-wall into the bell-cavity. This aboral pole is conical and terminates in a sucker-
shaped depression. There is a row of 5 to 7 short, somewhat club-shaped tentacles around
the oral pole and two closely-set rows of alternately arranged tentacles near the aboral pole.
Both aboral rows have together about 12 tentacles, and these are somewhat longer than the
5 or 7 oral tentacles. The middle part of the body of the actinula is devoid of tentacles.

FIG. 38. — Hydroid and medusa of Margelopsis harckelii, after Hartlaub, in Nor.iisdu-s Plankton.
Medusa is about to set free an actinula larva.

Medusa-buds develop after the actinula has been set free and has become about I mm.
long. These medusa-buds develop in clusters from the sides of the body of the actinula close
to the bases of the lower row of aboral tentacles with which they alternate in position. The
medusa-buds develop 2 or 3 tentacles in each of their 4 marginal clusters while still attached
to the actinula. Hartlaub found large numbers of these interesting hydroids floating in the
water at Helgoland, late in August.

Miiller, 1908, has studied the origin and structure of the ova in the medusa. In common
with HvbocoJon,tYie eggs are large, amoeboid, and few in number when full grown, for the
successful eggs devour the weaker ones in the ovary. The ooplasma is composed of wide
irregular fibers, forming a somewhat narrow-meshed network. The exoplasma is distinct
from the endoplasma and is wider in its meshes.

Margelopsis stylostoma Hartlaub.

Margelofiis slylostoma, HARTLAUB, 1903, Zoolog. Centralblatt, Bd. 10, p. 2S, fig. 2, fig. 4 ( ?)i 1904, Wisscn. Mccrvsuntrrsuch.
Kommis. Deutsch. Meere, Abth. Helgoland, None Folge, Bd. 5, p. 99, fig. I, fig. z (?); I9°7> Nordisches Plankton, Nr.
i*, p. 91, fig. 87 (88?).

This is a small spindle-shaped, free-floating hydranth, with a sucker-like expansion at
its aboral pole, and with three verticils of tentacles, i. e., 4 short-knobbed oral tentacles, a

7

82

MEDUSA OF THE WORLD.

middle zone of about 10 to 12 long-knobbed tentacles, and a basal (aboral) zone of" about 10
to 12 long-knobbed tentacles. Medusa-buds (?) This form closely resembles Tiarella
singularis F. E. Schulze, 1876, a small, isolated, attached hydroid found at Trieste. T ' . singu-
lans produces medusa at the base of the polypite below the third zone of tentacles.
Margelopsis stylostoma was found at Roscoff, northwest coast of France, in June.

FIG. 39. — Hydroid of Margelopsis stylostoma, after Hartlaub, in Nordisches Plankton.
FIG. 40. — Margelopsis hartlaubii, after Browne, in Bergens Museums Aarbog, 1903.

Margelopsis hartlaubii Browne.

Margelopsis hartlaubii, BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 10, plate i, fig. 2, plate 3, fig. 3. — HARTLAUB, 1907,
Nordisches Plankton, Nr. 12, p. 92, fig. 89.

Bell 2 mm. wide, 2 mm. high. Egg-shaped, with thick walls. 8 marginal tentacles, 2
upon each radially placed marginal bulb. These tentacles are ringed with nematocysts, and
are somewhat shorter than the bell-diameter. No ocelli. 4 narrow radial-canals. Stomach
large, conical, with a broad, flat, quadrangular base, without an axial, apical canal. Mouth
a simple, round opening. Gonad encircling the stomach. Color (in formalin): Gonads,
stomach, and tentacle-bulbs yellowish-brown. Three specimens; coast of Norway, Osterfjord
and Herlfjord, from depths of o to 200 fathoms.

Margelopsis gibbesi Hartlaub.
Plate 9, figs. 4-7.

Ncnwpsis gibbesii, McCfiADY, 1857, Gymn. Charleston Harbor, p. 163, plate 10, figs. 4-7. — FRECH, 1898, Lptli.ua pal<eozoica,
Theil i, Bd. I, Lfg. 3, p. 565, Stuttgart (compared with the Graptolites as a floating Tubularian).

Margelopsis gibbesi, HARTLAUB, 1903, Zoolog. Centralbl., Bd. 10, p. 28; 1899, Nachricht. kgl. Gesell. Wissen. math.-phys.
Klasse, Gottingen, Jahre 1899, p. 223, fig. 4.

Bell somewhat higher than a hemisphere, 2.5 mm. high, and walls quite thin and uniform.
There are 4 radially situated clusters of marginal tentacles. Each cluster contains 5 or 6
tentacles which arise from a large common basal bulb. Tentacles taper regularly from base
to tip, but the tip terminates in a knob of nematocysts. There are about 15 rings of nemato-
cysts upon the shaft of each tentacle and the entodermal axis of the tentacle is composed
of chordate cells resembling those of Obeha. There are 4 straight, narrow radial-canals.
Velum well developed. Manubrium wide and flask-shaped, and mouth a simple, round
opening without prominent lips, and without oral tentacles. In the female the ova project
prominently from the surface of the manubrium. The gonad surrounds the stomach on all
sides. The bell of this medusa is highly contractile, often drawing together so as to cause
the mouth to project beyond the velar opening. When the bell is relaxed, however, the manu-
brium extends but little more than half-way from the inner apex of the bell-cavity to the
velar opening.

PLATE 9.

Fig. I. Cladonema perkinsn. Nassau Harbor, Bahamas.

Fig. 2. Cladonema mayen. Moat of Fort Jefferson, Tortugas, Florida,

July, 1905.
Fig. 3. View looking down upon the aboral end of the stomach of ClaJont ma

mayeri, showing the arrangement of the radial-canals.
Fig. 4. Margelopsts gibbesn, male. Beaufort, South Carolina, December

24, 1904.
Fig. 5. Margelopsis gibbesn, male. Oregon Inlet, Pamlico Sound, North

Carolina, November 12, 1904.

Fig. 6. Margelopsis gibbesn, female. Cape Fear, North Carolina, Decem-
ber I, 1904.

Fig. 7. Tentacle of Margelopsis gibbesii.
Figs. 8 and 9. Stomotoca dinerna, young females. Agassiz Laboratory,

Newport, Rhode Island, July, 1892.
Fig. 10. Stomotoca dinema, young male. Agassiz Laboratory, Newport,

Rhode Island, Septemper 15, 1896.

Fig. I, copied from a drawing by Prof. Henry F. Perkins.
Figs. 2 to 10, from life, by the author.

PLATE 9

10

ANTHOMKnrS.E — MARGKLOPSI.S, l'KI.A( ;< iin I'll \.

83

The entoderm of the four large tentacle-bulbs is dull-yellow. There are no ocelli. The
entoderm of the manubrium was dull-\'ellow in two medusx, and dull-green in another. The

J O

difference is not a sexual one, however, for one ot the males was yellow while the other \\as
dull-green. The two yellow-colored medusa; were found, one in Oregon Inlet, Pamlico
Sound, North Carolina; and the other at Southport, Cape Fear, North Carolina, in Novi-mln-r.
The green-colored male (plate 9, fig. 4) was found in Beaufort River, at Beaufort, South
Carolina, on December 24, 1903, 12 miles from the ocean, in brackish water.

McCrady, 1857, presents a good figure of that which is probably the free-floating hydroid
of this species. The hydroid is pear-shaped with a hollow cup-like projection from its broad
aboral end. There are 2 whorls of tentacles, a circlet of about 8 tentacles around the broadest
zone near the aboral end and a circlet of about 6 tentacles near the mouth. A number of
medusa-buds develop from the sides of the hydranth in the middle region between the 2
circlets of tentacles. When set free the medusae have 4 short tentacles in each radial cluster.
A single specimen was found by McCrady in Charleston Harbor. The cup-shaped projection
from the aboral end of the hydranth is a feature which recalls the pneumatophore of the
Siphonophorae.

Genus PELAGOHYDRA Dendy.

Pelagohydra, DENDY, 1903, Quart. Journ. Microscop. Sci., vol. 46, p. i. — HARTLATB, 1903, Zool. Centralblatt, p. 27.

The type species is Pelagohydra mirabilis, Dendy, from New Zealand.

GENERIC CHARACTERS.

Medusa similar to Margelopsis. Hydroid pelagic and closely resembles that of Marge-
lopsis, but tentacles arise irregularly from the sides ot the hydranth, and are not arranged in

definite circlets as in Margelopsis,

Pelagohydra mirabilis Dendy.

Ptlagohyiira mirabilis, DENDY, 1903, Quart.
Journ. Microscop. Science, vol. 4(1, p. i.
plate 12. — HARTLAIB, 1903, Zool. Cen-
tralbl., p. 27, fig. 3.

Hydroid pelagic, with a trans-
lucent globular body 25 mm. wide,
constituting a float and a proboscis
at the end of which the mouth is
situated. The float bears many
long, irregularly arranged tenta-
cles. The proboscis is elongate,
pink in color, and of the shape of
a truncated cone. It has several
irregular whorls ot oral tentacles,
the shortest being nearest the
mouth. Medusae are borne upon
branching stolons which arise from
the sides of the float between the
tentacles. When about to be set
free the medusa is about i mm.
wide, with 4 radial groups of m.i i -
ginal tentacles, each group arising
from a large, pink, basal bulb, and consisting of 5 tentacles, 2 large, .2 small, and i smallest
median tentacle. 4 radial-canals. Manubrium pyriform and without oral tentacles. Nt\\
Zealand.

Family CLADONEMIDJE.

CladonemiJt, GKI;I:NBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 220. — HM.CKI.I.. 1879, Sys,t. ilcr Medusen, p. 98. — HART-
LAI-B, 1887, Zool. Anzcigcr, Bd. 10, p. 651; 1907, Nordisches Plankton, Nr. 12, pp. 5, no. — GUNTHER, 1903, Mittheil
Zool. Sta. Ncapel, Bd. 16, pp. 56, 57.

FIG. 40*3. — Pelagoh\dra mirabilis, after Dcmiy, in Quarterly Journal Micr
scop. Science.

a, entire li\ilr<>i,l, twice natural size.

b, enlarged view, showing medusa-buds developing among tentacles.

84 MEDUSA OF THE WORLD.

This family is of such interest that we devote more than the usual space to its consider-
ation. Haeckel, 1879, classifies it as a family coordinate with the Codonidae, but Hartlaub,
1887, showed that the genus Cladonema had an encircling gonad as in the Codonidae. The
medusa; are, however, highly specialized, both in respect to their anatomy and their mode of
life, and they are undoubtedly derived from some more simply-organized medusae among
the Codonidae. The hydroid of Cladonema is a StanriJia, and is closely related to Syncoryne,
the hydroid of Sarsia.

CHARACTERS OF THE FAMILY CLADONEMID^.

Anthomedusae with feathered or branched marginal tentacles, and with four or more
simple or branched radial-canals. The gonads may be ring-like and encircle the stomach, or
they may be more or less separated so as to be interradial or adradial, or developed in a spec-
ialized brood-pouch above the stomach.

Hartlaub, 1887, and Perkins, 1902, show that in the genus Cladonema the genital products
are developed over the entire gastric portion of the manubrium and the gonads are not con-
fined to restricted meridians, as was believed to be the case by Haeckel. Hartlaub found that in
Cladonema radiatum of the Mediterranean the genital products begin to develop in the ento-
derm. This species exhibits a successive hermaphroditism, although either sperm or ova may
precede. Perkins, however, in his study of Cladonema pcrkinsn of the Bahamas, found that
the genital products were developed in the ectoderm of the manubrium, and he failed to find
sperm or ova in the entoderm.

In Eleutheria, Hartlaub, 1 886, 1887 (Zool. Anzeiger), showed that the genital products
are developed exclusively in the ectodermal lining of a peculiar brood-pouch above the stomach,
and that the medusa is hermaphroditic. The brood-pouch is derived from the ectodermal
layer of the bell-cavity and is not connected with the stomach of the medusa, but communi-
cates with the bell-cavity by means of 6 interradial-canals.

Hartlaub, 1887, divides the Cladonemidae into two groups, one with and the other without
an apical brood-sac above the stomach. As this brood-sac is highly variable in its develop-
ment, some specimens of Eleutheria apparently failing to develop it and, moreover, as it is a
character which develops during the growth of the medusa and is not found in the young
animal, it would seem that Haeckel's classification of the Cladonemidae, based as it is upon
the possession of constant characters, is to be preferred. Moreover, as has been shown by
Giinther, 1903, we can not yet state that the brood-pouch of other genera of Cladonemidae is
similar in structure to that of Eleutheria.

As is well known, Haeckel, 1879 (Sitzungsber. Medicin. Naturwiss. Ges. Jena; see also
Chun, 1880, Ctenophoren des Golfes von Neapel), drew an ingenious comparison between
Ctenaria and the Ctenophorae, and concluded that the Ctenophorae might have been derived
from some form of Anthomedusa. Ctenaria appears to resemble the Ctenophorae in its 2
feathered tentacles, 4 bifurcated radial-canals, and in a peculiar sheath at the base of each
tentacle. However, Hartlaub's discovery that the brood-pouch in the closely-related Eleu-
theria is of ectodermal and not entodermal origin makes it appear that the resemblances
between Ctenaria and the Ctenophorae are of the nature of a mere parallelism and not indic-
ative of a genetic relationship. Haeckel compared the 8 adradial longitudinal lines of
nematocysts upon the exumbrella of Ctenaria with the 8 rows of ciliated combs in the
Ctenophorae. An even more remarkable case of convergence is shown by Kofoid, 1905
(Bull. Mus. Comp. Zool. at Harvard College, vol. 46, p. 163, i plate), in the case of the
Cystoflagellate, CraspeJotella, which is bell-shaped and provided with a well-developed velum,
the walls of which are contractile so that the mode of locomotion is similar to that of a medusa.
There is thus an external similarity of form between this protozoan and a medusa offering an
instance of convergence of a most striking character. Another remarkable instance of conver-
gence is that of the pelagic holothurian Pelagothuria natatrix, which bears a close resemblance
to a medusa (see Mem. Mus. Comp. Zool. at Harvard College, vol. 17, 1894). I believe also
that the bell of the Narcomedusa? is not homologous with that of the Anthomedusae and
Leptomedusae, but is a mere outgrowth from the walls of the actinula larva.

ANTHOMEDUS.E — ZANCLEA. 85

The Cladonemidae are close relatives of the Codonidae, this being illustrated most cleaily
by their hydroids, and they may properly be regarded as highly-specialized Codonidae, as has
been maintained by Hartlaub. For purposes of classification, however, I think it will be
well to permit them to remain in a family of Anthomedusae coordinate with the Codonidae, for
there is no single character which they have consistently in common with the Codonidae. For
example, the gonad in ClaJonemamzy be readily likened to that of the Codonidae, for it encircles
the stomach on all sides, but in some species of Zanclea the gonads show a decided tendency
to segregate interradially, and in Elcutheria they are developed in an ectodermal brood-sac
which is not connected with the manubrium.

The most constant character of the Cladonemidae is their forked, or feathered, marginal
tentacles.

The greatest difficulty in any attempt to classify the Cladonemidae among themselves
arises from our lack ot understanding of the nature of the brood-sac or apical cavity in the bell
ot the genera Pteroncma, Ctenarta, and Dendronema; and no classification worthy of serious
consideration can be effected until we know whether this sac is a mere extension of the stomach
or an ectodermal reproductive cavity similar to that studied by Hartlaub in Elcutheria.

FAMILY: CLADONEMIDAE. Subfamily Pteroneminai

Manubrium without oral tentacles. With 4 to 8 simple radial-canals.

Zanclea GEGENRAUR, 1856. With 2 to 4 marginal tentacles at bases of 4 radial-canals. The tentacles give rise to an
abaxial row of simple branches, each terminating in a cluster of ncmatocysts. With meridional rows of nema-
tocysts upon the exumbrella. No ocelli. No brood-pouch above the stomach. The hydroid is Gtinmaria McCrady,
Allman.

Zancleofish HARTLAUB, 1907. Similar to Zanclea, but with tentacular ocelli and without meridional rows of nemato-
cysts.

Pteronema HAECKEL, 1879. With 4 well-developed tentacles similar in structure to those of Zanclea. With 4 simple
radial-canals. With a brood-sac above the stomach. Hydroid unknown.

Eleutheria QTATREFAGES, 1842. With 4 to 6 simple radial-canals, and an equal number of bifurcated tentacles. With
a brood-pouch above the stomach. This brood-pouch is not connected with stomach-cavity, but communicates
\vith bell-cavitv by means of simple interradial pores. Genital products develop exclusively in brood-pouch. Medusa
is hermaphroditic. Hydroid is Clavatella Hincks.

Mnestra KROHN, 1853 (Archiv. fur Naturges., Jahrg. 19, Bd. i, p. 278). Medusa degenerate, sessile upon Pliyllirhoc
in the Mediterranean. With 4 to o small marginal tentacles having nettling capsules along their abaxial sides. A
ring of nematocysts extends around the bell-margin, and 4 linear tracts of nettle-cells extend upward over the
exumbrella, t from each tentacle-bulb. With 4 simple, unbranched radial-canals and ring-canal. No brood-pouch
above stomach. Throat of medusa blocked by a spongy mass of entoderm. Medusa attaches itself by its mouth to
throat of Phyllirhoe, and the larvae of the Mnestra probably develop within the gastric cavity of the Phyllirhoe* It
is elaborately described by Gunther, 1903 (Mitt. Zool. Sta. Neapel, Bd 16, pp. 35-62, plates 2, 3).
FAMILY: CLADONEMID.'E. Subfamily Dendroneminte:

The mouth is surrounded by simple, or branched, oral tentacles. Some or all of the radial-canals bifurcate.

Ctrnaria HAECKEL, 1879. With 2 marginal tentacles, each of which gives rise to a row of nematocyst-bearing filaments.
With 4 bifurcated radial-canals. With simple, unbranched, oral tentacles. With an apical sac above the stomach.
Hydroid unknown.

Clailonema DTJARDIN, 1843. With 4 to 5 bifurcated radial-canals, or 8 to 10 simple radial-canals, or with both simple
and bifurcated radial-canals. With a tentacle at foot of each radial-canal. These tentacles branch dichotomouslv
or complexly, and branches end in suckers or nematocyst-knobs. With simple, unbranched, oral tentacles. No
brood-sac above stomach. Hydroid, Siauridia Dujardin.

Deridroneiiia HAECKEL, 1879. With bifurcated radial-canals, and dichotomously branched tentacles. Branches end in
suckers, or in nematocyst-knobs. Mouth surrounded by 4 groups of dichotomously branched oral tentacles. There
is an apical sac above the stomach.

Genus ZANCLEA Gegenbaur, 1856.

Zanclea, GEGENBAVR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 229. — KEFERSTEIN UNO EHLERS, 1861, Zool. Beitr. in Xt-apcl und

Messina, p. 85. — AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344.
Zariflea-t- Gemtntiriii, HAE( KEI., 1879, Syst. der Mcdlisen, pp. IO2, 103.
Zanclea, FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 150. — HARTLAI/B, 1887, Zoo], Anzeiger, Bd. 10, p. 654; njo^, N'or-

dischcs Plankton, Nr. 12, p. 112. — VANHOFFEN, 1891, Zool. Anzeiger, Jahrg. 14, p. 445. <!i NIH>K. i'|o}. Mitt. Zool.

Sta. Neapel, Bd. 16, p. 57.

Gemmaria= Zanclea, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 750.
Gemmaria, McCfiADY, 1857, Gymn. Charleston Harbor, p. 49. — A<:», i '. \ . iSi^, North Amcr. Acal., p. 184. An \nx, 1872,

Monog. Tubul. Hvdroids, p. 289. — BROWNE, 1896, Proc. X.ool. Sot. London, p. 4111.

\'on Gemmaria, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard I'. .Hi vol. 8, p. 150. •

Gemmaria, HARTLAI'B, 1887, Zool. Anzeiger, Bd. 10, p. 6^4. -fJiM'Hm, [903, Mitt. Zool. Sta. Neapel, Bd. if>, p. 57.

Gemmaria (hydroid), HARTLAI R, 190$, Zoologischen Jahrbiich., Supplement 6, p. 527.

Corvnith, Mi r. i:v 11. 1^90., Quart. Journ. M ici., new ser., vol. 42. p. ^4. \t TTING, 1901, Bull. I'. S. Fish Commission

for iScfij, pp. ^29, 372. — HARGITT, 1904, Bull. U. S. Bureau uf Fisheries, vol. 24, p. 42.
Halorharis (hydroid), Ai.tssiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 239, plate 20, figs. io-ioc.
Gemmaria+ Zanclea, HARGITT, 1908, Biol. Bulletin, vol. 14, p. 104.

The type species of this genus is Zanclea costata Gegenbaur, 1856, of the Mediterranean.

Mi

MEDUSAE OF THE WORLD.

Tabular Description of the Medusa of Zanclea.

Z. costata Gegenbaur,
1856.

Z. gemmosa McCrady,
1857-

Z. implexa Allman,i864

Z. cladophora = Gem-
maria cladophora A.
Agassiz, 1865.

Shape and size of bell
in mm.

Ellipsoidal. 4 to 5 high,
3 to 4 wide.

Pyriform. 6 high, 4
wide.

Dome-like to globular.
1.5 high, 1.75 wide.

Pyriform. Size (?)

Length of the 4 merid-
ional rows of nettle-
cells above tentacle-
bulbs, in terms of

From tentacle-bulbs to
very near a pex of bell .

One-fourth to one-third
distance from margin
to apex.

0.25 to 0.5

°-5

bell-height.

Number and length of
tentacles in terms of
bell-radius (r).

4 tentacles, each 3 to 4 r
long-

4 tentacles, each 3 to 4 r
long.

2 to 4 tentacles, each
about 4 r long.

2 long and 2 short ten-
tacles; long tentacles
4 r, short over one-third
r long.

Character of abaxial
filaments upon ten-
tacles.

Very numerous, slender,
and ending in a spheri-
cal capsule containing
3 to 5 nematocysts.

Very numerous, slender,
and ending in a spheri-
cal capsule containing
3 to 9 nematocysts.

Very numerous, slender
filaments, each ending
in an oval capsule
containing 4 to 8
nematocysts.

Very numerous, slender
filaments, ending in
knob-like clusters of
nematocysts. Num-
ber of nematocysts in
each knob ?

Length and shape of
manubrium.

Not quite half as long as
depth of bell-cavity.
Flask-shaped.

Flask-shaped, 4-sided.
As long as depth of
bell-cavity . Mouth
with 4 small lips.

Urn-shaped. Only half
as long as depth of
bell-cavity. Mounted
upon short peduncle.
Mouth round opening.

Conical. 4 prominent
lips.

Gonads.

4 interradial masses of
eggs on swollen sides of
stomach.

On 4 interradial sides of
stomach. Much
swollen.

On upper sides of
stomach, more or less
separated in 4 chief
radii.

Gonad encircles stom-
ach.

Color.

Gonads red, tentacles
yellowish-brown.

Entoderm of stomach
and tentacle-bulbs
creamy-pink. Ento-
derm of manubrium
delicate green.

Stomach and tentacle-
bulbs brownish-red to
crimson. Entoderm
of tentacles orange.

Tentacles light-brown.
Orange and yellow
spots in tentacle-
bulbs. Color of stom-
ach ( ?)

Where found.

Mediterranean. Common.
Medusa having only 2
tentacles, but in other
respects similar to Z.
costata found at Tor-

West Indies northward
to Woods Hole.

Atlantic coasts of west-
ern Europe to Medi-
terranean.

Coast of Massachusetts,
Nahant, Woods Hole.

tugas, Florida.

Hydroid.

Hydroid ( ?) (see text).

Hydroid: Gemmaria
gemmosa.

Hydroid: Gemmaria
implexa.

Hydroid unknown.

GENERIC CHARACTERS.

Cladonemidae with 4 simple radial-canals, and 2 to 4 radially situated tentacles which give
rise to filiform, knobbed, nematocyst-bearing side branches. The gonad is developed upon the
sides of the stomach. There is no apical brood-pouch above the stomach. There are merid-
ional lines of nettle-cells over the exumbrella. There are no tentacular ocelli. The hydroid
is Gemmaria Allman, 1871.

In 1857, McCrady founded the genus Gemmaria for a Zanclea-\ike medusa with only 2
tentacles. Vanhoffen, 1891, and Browne, 1905, have observed that the 2-tentacled young of
other species of these medusae often becomes 4-tentacled when adult, and according to Browne,
1905, "Gemmaria" implexa may become mature with 2 or with 4 tentacles. It is evident,
therefore, that there are no definite generic distinctions between Zanclea and Gemmaria, the
"Gemmaria" condition being only a developmental or retarded phase of Zanclea.

Murbach and Nutting have erroneously applied the name "Corynitis" to a 2-tentacled
medusa of Zanclea. Apparently these authors had not seen the medusa of Corynitis, which is

ANTHO.MKDCS.E — ZANCI.KA.

well figured by McCrady. I have frequently taken Corynitis in Charleston Harbor, South
Carolina, where McCrady discovered it, and it is wholly different from any "Geniniaria" or
Zanclea. (See also Hargitt, 1908.)

The common '/aticli-a ot the southern coast of New England, in America, may become
mature with only 2 tentacles, and with poorly developed lips. On the other hand, a vcrv
similar medusa is occasionally found in the same waters with 4 tentacles and with 4 well-
developed lips. I am inclined to believe that these conditions represent developmental staues.
or starved and well-fed states of one and the same medusa.

Zanclea costata Gegenbaur.
Plate 8, figs, z, 3, 6, and 7.

Zanclea conaia, GEGENBAUR, 1856, Zeit. fur wisscn. Zool., Bd. 8, p. 229, taf. 8, figs. 4-7. — HAECKEL, 1879, Syst. dcr Mcdu sen

p. 103.
Hydroid ( ?), MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 35, plate 41, figs. 137, 138.

Gegenbaur found many specimens of this medusa in the Mediterranean. His fully

developed medusae had 4 long, equally developed tentacles. For details see tabular description

of the medusae of Zanclea.

I have found a medusa at Tortugas, Florida, which accords well with 7, ancle a costata,

excepting that it has but 2 tentacles.

Medusa from Tortugas, Florida. — Bell 6 mm. high and ellipsoidal, with bluntly-rounded

apex. Without an apical projection. Bell-walls of uniform thickness everywhere. There are

2 long, spindle-shaped, rough-looking tentacles somewhat longer
than the bell-height. These are thickly covered with papillae and
beset on their outer sides with a row of filaments, each of which
ends in a spherical knob, the surface of which is covered with
blunt papillae. Each of these knobs contains a cluster of nemato-
cysts (plate 8). Basal bulbs of tentacles are elongate, spindle-
shaped, and hollow. No traces of tentacle-bulbs 90° away from
the 2 long tentacles. 4 swollen lines ot nematocysts extend upuaul
over the exumbrella from the bases of the 4 radial-canals and
nearly meet near the apex of the bell. Velum well developed. 4
radial-canals, and the circular vessel straight and narrow. Manu-
brium flask-shaped, with 4 simple lips. Entoderm of the manu-
brium and tentacles milky white, There were 4 interradial
gonads. A single specimen was found in a surface-tow at Tor-
tugas, Florida, on May 15, 1906.

This species is distinguished by its ellipsoidal, high, rounded
bell, without a sharp-pointed apex, the 4 very long meridional lines
of nematocysts over the exumbrella, and the spherical nematooyst-
capsules at the ends ot the tentacular filaments.

The Tortugas medusa differs from //uiclea cost<ita,as described
by Gegenbaur, in having but 2 tentacles. It is well known, hu\v-

Flc. 41. — Zanclea costata, from the
Mediterranean, after Gegen-
baur, in Zeit. fiir wissen.
Zool., Bd. 8.

ever, that in Zanclea some individuals ot the same species develop 4 while others become
mature with but 2 tentacles.

Gegenbaur states that the female gonads are red (pink '). In the Tortugas specimen the
male gonads were milky-yellowish white.

Hydroid (plate 8, fig. 2). — That which is probably the hydroid stock of this species was
tound by Mayer, 1900, at Tortugas, Florida, and erroneously ascribed to '/.anclea gctntnosa;
but Murbach, 1899 (Quart. Journ. Micros. Sci., New Series, vol. 42, p. 354), describes and
figures the hydroid ot '/.anclea genniiosa, and it is now evident that the form described In
Mayer is specifically distinct from Z. gemmosa and is probably Zanclea costata, the medusa
of which occurs at Tortugas.

Hydroid of Zanclea costata (?). — The hydrorhiza is creeping, tubular, and net-like, and
gives rise at irregular intervals to short, more or less twisted hydrocauli. Both the hydrorhiza
and hydrocauli are covered with a horny, chitmous pensarc, which is annulatcd in the hydro-

MEDUSAE OF THE WORLD.

caulus. The hydrocaulus is corrugated and opaque in color throughout its length, and in
this respect differs from the European G. nnplcxa Allman, 1871. The fully developed
hydranths are only 1.5 mm. in height. They are elongate, spindle-shaped, and taper gradually
from near the base to the summit, so that the diameter near the mouth is less than that near
the base. The tentacles arise in 5 to 8 well-defined whorls from sides of hydranth, each whorl
consisting of 4 to 6 tentacles. These tentacles are short and taper from base to end, termi-
nating in a large knob containing nematocysts. The entodermal cores of the tentacles are
composed of vacuolated cells. The tentacles are stiff and project rigidly outward from the
sides of the hydranth. 4 to 8 medusa-buds in various stages of development arise from the
sides of the hydranth near the base immediately below the lowest whorl of tentacles.

When set free the young medusa (plate 8, fig. 3) has 2 well-developed, radially situated,
diametrically opposed tentacles, and 2 rudimentary tentacle-bulbs. There are 4 radially
situated, nematocyst-bearing ridges upon the exumbrella, and these extend about half-way
up the sides of the bell. The bell-walls are very thin and flexible. There are 4 straight,
narrow radial-canals, and the manubrium is a short, simple tube with no trace of the gonad.
The mouth is a simple, round opening. Before being set free the tentacles are coiled inward
so as to lie within the bell-cavity. Soon after the medusa has become liberated, however,
the tentacles uncoil and project outward. The deep-lying entoderm of the hydranths is a
delicate creamy-pink, while the more superficial entoderm is translucent and milky in color.
The cells of the superficial entoderm are large and vacuolated. The hydrorhiza and hydro-
caulus is protected by a horny, yellow, amber-colored, chitinous sheath, but this does not
extend over the hydranths. The hydroid was found at Tortugas, Florida, in June, 1899,
growing upon a piece of floating gulfweed (Sargassum).

Zanclea gemmosa McCrady.

Plate 6, fig. 7; plate 7, fig. 5; plate 8, figs. 4 and 5.

Gemmaria gemmosa, McCRADV, 1857, Gymn. Charleston Harbor, p. 49.

Zanclea gemmosa, McCRADV, 1857, Gymn. Charleston Harbor, p. 49, plate 8, figs. 4, 5. — AGASSIZ, L., 1862, Cont. Nat. Hist.

U. S., vol. 4, p. 344. — HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 121, fign. 116, 117.
Gemmaria gemmosa, AGASSIZ, A., 1865, North Amer. Acal., p. 184, fig. 306. — HAECKEL, 1879, Syst. dcr Medusen, p. 104. —

FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, No. 8, p. 150, plate i, figs. 8-12. — FEWKES, 1884,

American Naturalist, vol. 18, p. 196, fig. 2.
Corynitis agassizii (hydroid), MURBACH, 1899, Quart. Journ. Micr. Sci., vol. 42, p. 354, plate 34, fig. 12 (medusa, figs, n,

I3» H-)

Gemmaria sagiltaria, HAECKEL, 1879, Syst. der Medusen, p. 103, taf. 7, figs. 3, 4 (young medusa).
Zanclea gemmosa (hydroid), WALLACE, 1908, Carnegie Institution Year Book, No. 7.

Adult medusa. — Bell 6 mm. in height, and pynform. There is a well-developed, solid,
apical projection. Side walls quite thin. There are 4 long, radially situated tentacles with
large, hollow basal bulbs, which lack ocelli. The shafts of the tentacles are crowded with
large numbers of short filiform branches, which arise only from the dorsal (exumbrella)
side of the tentacles, although the twisting of the tentacle may cause them to appear to arise
from all sides. Each of these filiform branches terminates in a large spear-head-shaped
body which bears a number ot long, delicate, sharp-pointed bristles, and which contains
3 to 9 egg-shaped nematocyst-cells. A club-shaped swelling extends longitudinally from
the base of each tentacle a short distance up the side of the exumbrella. These swellings are
thickly covered with nematocysts. There are 4 straight, slender radial-canals, and a narrow
circular vessel. The velum is wide and delicate. The manubrium is flask-shaped, and cruci-
form in cross-section and extends to the level of the velar-opening. The mouth is a simple,
cruciform opening without prominent lips. The gonad is upon the sides of the stomach.
These sides are greatly swollen and give to the manubrium its flask-shaped appearance.
The entoderm of the manubrium and tentacles is of a delicate creamy-pink and the ectoderm
of the manubrium is of a delicate shade of green.

Young medusa (plate 7, fig. 5; plate 8, figs. 4, 5). — In the medusa, I mm. in height,
there are but 2 marginal tentacles. The bell-walls are very thin and the manubrium is a
simple fusiform tube. There is no apical projection to the bell, and no trace of gonads upon
the manubrium, and the mouth is a simple, round opening without cruciform lips.

This species extends from the West Indies to the southern coast of New England. It is
quite common in Newport Harbor late in the summer.

ANTHOMEDUS^E — ZANCLKA.

S'.)

The hydroid stock has heen described by Murbach under the name of "Cor\nitis
agiissizii," but is evidently a Gtmnmria. The hydranths are 1.5 to 2 mm. in length and
arise singly from a slender, thread-
like, anastomosing hydrorhiza. The
hydrorhiza is covered with a dc-h-
cate perisarc, but this does not
extend over any part of the hy
dranths. In this respect the species
is very different from Z. costiiin.
The hydranths are also more cylin-
drical and uniform in width through-
out the polypite than in Z. fostatn.
The tentacles are irregularly ar-
ranged and the medusa-buds are
found not only near the base of the
hydranth, but scattered along the
sides. In Zanclea costata they are
confined to a zone immediately under
the lowest circlet of tentacles. The
color of both forms is quite similar,
excepting that the hydrorhiza is deep
pink in the Z. gemmosa and rosin-
yellow intheZ.co.sta/rt. W.S.Wallace
found this hydroid growing upon
floating Sargassum at Tortugas,
Florida, in July, 1908.

The young medusa
here figured was obtained

at Newport, Rhode Is- Fic.42.— Z0nr/fagfmmo5a,fromFewkes,inBull.Mus.,Comp.Zool. at Harvard College.

land, and was identified by Dr. Alexander Agassiz as the young of '/.anclca gemmosa.
Z. gemmosa readily devours the medusae of Obeha.

Zanclea iraplexa Allman.

Cor\nt pelAgiea+ Corytif implexa (hydroid and medusa), ALDER, 1857, Catalogue Zoophytes Northumberland, Trans. Tync-

sidc Nat. Field Club, vol. 3, pp. 13, 18 (of the "separate"), plate 7, figs. 1-6.
Zanclea implexa, ALLMAN, 1864, Annals and Mag. Nat. Hist., vol. 13, p. 357. — HINCKS, 1868, Hist. British H^lr.n.l /.. ',,)..

p. 59, plate 9, fig. 3. — HARTLAUR, 1907, Nordisches Plankton, Nr. 12, p. 116, fign. 106-111 (list of localities, authors,

and best description of the medusa).
Gemmariii implexa, ALLMAN, 1871-72, Monog. Tubularian Hydroids, pp. 223, 290, plate 7, figs. i-io. H \M x n . iX-<>. V. i.

dcr Medusen. p. 105. — JICKELI, 1883, Morphol. Jahrbuch, Bd. 8, p. 6n, taf. 26 (hi-riiWv ). CKJ, N-.4. Arbeit

Zool. In.st. \Vien, Bd. 5, p. 353. — BROWNE, 1896, Proc. Zool. Soc. London, p. 461.— HARI.ITT, 1904. Mnth. /.nol. Si.iiinn

Neapel, Bd. 16, p. 574, taf. 22, figs. 27-29 (hydroid and medusa from the Bay of Naples). BKOWNI, 1905, Proc. Roy.

Soc. Edinburgh, vol. 25, p. 750.

This medusa is closely related to /.unclea gemmosa of the Atlantic coast of the I'mted
States, but it is much more swollen than the American form, and differs in other n-spivts.

The bell is globular with fairly thin side walls and a thick, solid, dome-like apex. It
is 1.5 mm. high, 1.75 mm. wide. There are 4 prominent nematoryst tracts extending one-
fourth to half the distance from the tentacle-bulbs up the sides of the bell, and the nettle-
cells in each tract are arranged in two rows.

There are usually 2 long tentacles 1 80° apart alternating with 2 short tentacle-bulbs,
but according to Browne, 1905, the medusa may become mature with 2 or with 4 long ten-
tacles. The long tentacles are about twice as long as the bell-diameter and their upper,
abaxial sides bear each a double row of numerous filaments, which terminate in oval capsules
containing 4 to 8 nematocvsts. The basal bulbs of the tentacles are globular, and lack ocelli.

There is a slight linear (glandular?) swelling occupying the middle third of each of the
4 radial-canals. The stomach is mounted upon a short peduncle and is urn-shaped, with
a simple, round mouth-opening. The gonads surround the stomach and are more or less

90

MEDUS/E OF THE WORLD.

separated in the 4 principal radii. The basal bulbs of the tentacles and the stomach are
brownish-red to bright crimson, and the entoderm of the tentacles is orange.

This medusa is found off the Atlantic coasts of Western Europe, and in the Mediter-
ranean. Graeffe, 1884, states that at Trieste, Adriatic Sea, the medusae bud forth from the
hydroids in the spring, and a second brood appears to develop in summer.

The hydroid is Gemmaria implexa, well described and figured by Allman, 1871-72.
The stems of the hydroid are erect, 6 to 8 mm. high, and arise at intervals from a creeping,

net-like hydrorhiza which is attached to
shells, stones, etc., in shallow water. The
stems are usually single, but occasionally
they branch. The perisarc covers the
hydrorhiza and extends as a transversely
corrugated, opaque tube up the lower
part of each hydranth. Below the
zones of tentacles the perisarc becomes
abruptly smooth, and ends in a slightly
flaring cup. The hydranths are cylin-
drical to club-shaped, with dome-like
peristome. There are 40 to 50 short
tentacles, each ending in a knob of
nematocysts. These tentacles are ar-
ranged in 10 to 13 irregular verticils of
about 4 tentacles each, the tentacles
being about 90° apart. The medusa-
buds are borne upon peduncles which
arise in clusters from the sides of the
hydranth below the middle, but above
the lowermost verticils of tentacles.

When set free the medusa has 2
well-developed tentacles which give
rise to numerous filaments, each ending
in a capsule bearing terminal bristles
and containing several nematocysts.
There are also 2 small tentacle-bulbs
90° apart from the well-developed ten-
tacles. The bell is oval and higher
than a hemisphere and 4 lines of
nematocysts extend up about one-fourth
to one-third of the bell-height, one above
each tentacle-bulb. The 4 radial-canals
are straight and simple, and the manu-
brium is a simple tube with a round
mouth-opening and straight cylindrical

Fie. 43.— Hydroid of Zanclea implexa, after Allman, in Ray sides. It lacks a peduncle and is about

Society, 1871-72. See also, Fig. 44. nalf as long as depth of bell-cavity.

The hydranths are white with pale-pink entoderm. The perisarc on the proximal portion
of the hydrocaulus is brown and on the distal portion, colorless. It is closely related to the
hydroid of Zanclea gemmosa.

Zanclea cladophora Hartlaub.

Gemmaria cladophora, AGASSIZ, A., 1865, North Amer. Acal., p. 184, figs. 307-310. — HAECKEL, 1879, Syst. der Medusen, p. 104. —
NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 371, fig. 78. — HARGITT, 1904, Bull. U. S. Bureau of

I-' i .lirrn-s, vol. 24, p. 42.
Zanclea cladophora, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 121, fign. 112-115.

Bell pyriform and about as broad as high. Apical projection solid, rounded, and dome-
like. There are 2 long, diametrically opposed tentacles from the outer sides of which arise
a great number of slender filaments, each terminating in a knob-like body which contains

ANTHOMEDUS.E — ZANCLKA. XAM MXH'SIS.

91

nematocysts. These long tentacles are 1.5 times as long as the bell-diameter. There are also
2 small, conical tentacles 90° apart from the long ones. The small tentacles do not give rise
to side-filaments. No ocelli. A ridge of nematocyst-cells extends from the base ot each ten-
tacle about half-way up the side of the exumbrella. Velum broad. There arc 4 broad radial-
canals and a narrow circular tube. The outline of the bell-cavity does not follow that of
the outer surface of the bell, but is widest at the inner apex. Manubrium conical, with a wide-
proximal base and regularly-narrowing sides. It does not extend quite to the level ot the
velar opening. The 4 lips are quite prominent, and the mouth is a rectangular opennm.
The mouth is surrounded by 2 or 3 rows of large nematocyst-cells, similar to those found
upon the sides of the exumbrella. The gonad is developed upon the sides of the stomach,
which it encircles. The tentacles are light-brown in color, and the entoderm at their bases
contains orange pigment. There are bright-yellow pigment-spots at the bases ot the two
rudimentary tentacles. This species was found at Xahant, Massachusetts, in 1862, by Dr.

Alexander Agassiz. Nutting, 1901, obtained
it at Woods Hole, Massachusetts, in August,
1899. Neither of these observers states the
dimensions of the medusa.

The peculiar outline ot the subumbrella
and the constriction above the lips figured
by A. Agassiz may be due to unnatural
contraction.

Many of the characters of this so-called
remain in doubt.

species

Zanclea nitida.

Gtmmaria nitida, HARTLAI B, 1905, Zoolog. Jahrbiichrrn.
Suppl. 6, p. 527, fign. J, H (hydroid).

PolvpitesaboutS mm. long. Club-shaped

rio. 44. — Zanclea ini[>lc\a, after Hartlaub, in Norillschcs rlankton. - ' .

to cylindrical, with about 50 short knobbed

tentacles, irregularly distributed. Hypostome short and flatly conical. Each polvpite IK i
i to 3 clusters of small medusa-buds. These are borne upon short branching stems which
arise between the tentacles near the lower, basal part of the polypite. The condition of the
medusa is not described. Found at Juan Fernandez Island, off the Pacific coast of South
America.

Genus ZANCLEOPSIS Hartlaub, 1907.
ZancleopaSf HARTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 115, 116.

GENERIC CHARACTERS.

Cladonemidae related to Zandca, but with large, stout, lateral branches on the tentacles.
With ectodermal ocelli upon the tentacle-bulbs, and without meridional clusters of nemato-
cysts upon the exumbrella.

The type species, and only known form, is Zancleopsis Jichotorna, from Tortugas, Florida,
described by Mayer under the name Gemimirni Jn-hotvma.

This genus is distinguished from Zatiflra by the large lateral branches upon its tentacles,
and by the absence of nettle-cell tracts upon the exumbrella. Also, it has <j> < Hi, and these
are not known in Y.nnclea.

Zancleopsis dichotoma Hartlaub.
Plate 8, fig. i.

Gtmmaria dichotoma, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 35, plate 17, fig. 40.
Zancleoftsis dichotoma, H^RTLArn, 1907, Nordisches Plankton, Nr. 12, p. 115, fig. 105.

Bell miter-shaped, with thin walls, and solid apical projection. 3 mm. high and about
2.5 mm. wide. There are 2 rudimentary tentacle-bulbs and 2 well-developed, diametrically
opposed tentacles. These long tentacles are of unequal length, one being about as long as

92 MEDUSA OF THE WORLD.

the bell-height, while the other is only about two-thirds this length. Each of these tentacles
terminates in a club-shaped, nematocyst-beanng end. The small tentacle gives rise to 2 to 3
side branches while the large one gives rise to 4 side branches. These side branches arise
from the outer (aboral) side of the tentacle, and each one terminates in a knob-like end con-
taining nematocysts as in the end of the main shaft itself. The youngest and least-developed
side branch is always found nearest the side of the bell, whereas the oldest is nearest the outer
end of the tentacle. The basal bulbs of the tentacles are large and swollen and contain each
an ectodermal ocellus upon the outer (abaxial) side of the bulb. The ectodermal core of each
tentacle and of the side branches is hollow. The nematocyst bulbs of the tentacles are
provided with delicate bristles. The velum is quite narrow. There are 4 straight, simple
radial-canals and a narrow circular vessel. The manubrium is flask-shaped, and cruciform in
cross-section, with 4 small, cruciform, nematocyst-covered lips. The stomach-cavity does
not project upward into the gelatinous apex of the bell. The gonads are developed upon
the interradial sides of the stomach.

The entoderm of the manubrium and the circular and radial-canals is dull ocher-yellow,
while the entodermal cores of the terminal bulbs of the tentacles is of a brighter yellow.

The 4 ocelli (2 on the bulbs of the well-developed tentacles and 2 on the rudimentary
tentacle-bulbs) are dark reddish-brown.

This medusa is occasionally found at Tortugas, Florida, from May until July. It is
taken each year upon the surface, but never in large numbers.

This species is widely separated from other medusae of the genus Gemmana by its well-
developed ectodermal ocelli upon the 4 tentacle-bulbs and by the total absence of nematocyst
tracts upon the exumbrella. The "filaments" arising from the abaxial sides of the 2 long
tentacles are almost as thick as the main shaft of the tentacle itselt, and in comparison with
species of Gemmaria they are greatly reduced in number. This numerical disparity is, how-
ever, counterbalanced by their great size.

Genus PTERONEMA Haeckel, 1879.

Microstoma, preoccupied by CUVIER, 1817, for Fishes.

Microstoma, LESSON, 1829, Voyage autour du monde sur la Coquille, tome 2, Zooph., p. 130.

Pteronema, HAECKEL, 1879, Syst. der Medusen, p. 101. — VANHOFFEN, 1891, Zool. Anzeiger, Jahrg. 14, p. 445.

The first medusa to be described which belongs to this genus is Microstoma ambigua
Lesson, from New Guinea.

Haeckel, 1879, describes that which is probably the same medusa, in an expanded state,
under the name Pteronema darwinii. This species is described from the coast of Australia,
but the exact locality is not stated.

GENERIC CHARACTERS.

Similar to Zanclea, but distinguished by having a brood-sac above the stomach, the
nature of which is unknown. If it be similar to that of Eleutlieria it has no connection with
the stomach. Haeckel considers it to be continuous with the stomach, but his studies do
not demonstrate this to be true, for he cut no sections.

There are no meridional tracts of nematocysts upon the exumbrella in Pteronema.

Pteronema darwinii Haeckel.

( ? ?) Microsioma ambigua, LESSON, 1830, Voyage de la Coquille, Zooph., tome 2, p. 130, plate 14, figs. 5, 5'.

(? ?) Microslotna ambiguus, LESSON, 1843, Hist. Zooph. Acal., p. 295.

(? ?) Zanclea ambigua, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344.

Pteronema darwinii, HAECKEL, 1879, Syst. der Medusen, p. IOI, taf. 7, figs. I, 2.

(? ?) Pieronema ambiguum, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 144 (literature, 1830-1850).

Haeckel describes his Pteronema darwinii from a single specimen preserved in glycerin.
He states that the preservation was good, but that the tentacles were contracted and are
represented in his figure as being of double the length observed in the preserved specimen.
This medusa was obtained somewhere ofF the coast of Australia.

Lesson, 1830, describes a medusa under the name Microstoma ambigua from Waigion,
New Guinea, which may possibly be identical with P. darwinii. Lesson's description is,

ANTHOMEDUS.E — FTERONEMA, ELEUTHEHIA.

93

however, too vague to be of service, and his figure 5' is almost equally hopeless. The spherical
bell is apparently turned inside out, and there are 4 short, feathered, marginal tentacles and
a flaring, conical manubrium, the tentacles and manubrium being bright yellow. His figure
5 is even worse for purposes of identification. I advise the dropping of Lesson's medusa
from future lists, for it will doubtless be impossible to determine it even generically.

In Haeckel's medusa the bell is 6 mm. high, 4 mm. wide, pyriform with a pointed

conical apex. There are no longitudinal lines of nemato-
cysts over the exumbrella. There are 4 radially placed
marginal tentacles, longer than the bell-height, and each
provided with an abaxial row of filiform side branches,
each one of which terminates in a spear-head-shaped
capsule containing 5 to 10 nettle-cells. The basal bulbs
of the tentacles are conical and lack ocelli. There are 4
straight, wide, jagged edged radial-canals and a narrower,
smooth edged circular vessel. The stomach is spindle-
shaped, halt to two-thirds as long as the depth of the
bell-cavity, and provided with 4 simple lips. Gonads ( ?)
on the adradial sides of the stomach.

The distinguishing characteristic of this medusa is
the elongate, spindle-shaped brood-pouch above the
stomach in the gelatinous apex of the bell. Haeckel
observed ripe eggs and gastrulae in this brood-pouch.
The anatomical relationships between this brood-pouch
and the stomach are unknown, and it may have no con-
nection with the stomach, but be ectodermal and con-
nected with the subumbrella epithelium as it is in
Eleutheria, Haeckel's description affords no solution of
this problem. Color ( ?) Coast of Australia. Exact local-
ity unknown. A single specimen described by Haeckel.

Genus ELEUTHERIA Quatrefages, 1842.

Elcufhcriti, (Ji'ATRKFAr.Es, 1842, Compt. rend. Acad. Set., Paris, tome 15, p.
168; 1842, Annal. des Sci. Nat., serie 2, tome 18, p. 270.

Clavatella (hydroid and medusa), HINCKS, 1861, Annals ami Mai:. Nat. Hist.,
scr. 3, vol. 7, p. 73.

Elfutheria, KROHN, 1861, Wiegemann's Archiv. fur Natures., Jalirj;. 2-, Bd.
i, p. 157. — CLAPAREDE, 1863, Beobacht. iiber wirbrllose ThiYre, p. 4. —
ALLMAN, 1872. Monog. Tubul. Hydroids, p. 384. — HAECKEL, 1879,
Syst. der Medusen, p. io<;. — HARTI-AVK, 1886, Zool. Anzeiger, Bd. 9,
p. 707; Ibid., 1887, Bd. 10, p. 652; Iliiil., Bd. 12, p. 665, 1889; 1907,
NorJisches Plankton, Nr. 12, p. 126. — BROWNE, 1902, Annals and
Mag. Nat. Hist., ser. 7, vol. 9, p. 279.— GUNTHKR, 1903, Mitt. Zool.
Sta. Neapel, Bd. 16, p. 57. — KRVMBACH, 1907, Zool. Anzeiger, Bd.
3 1, p. 450.

FIG. 45. — Ptcronema danvinii, after
Haeckel, 1879.

The type species of this genus is the extremely v.i li-
able Elfutheria dichotoma of the Mediterranean and
Atlantic coasts of Europe. Haeckel, 1879, p. 106, records 12 subspecies of this medusa. Tin-
best description is by Hartlaub, 1886, 1907.

GKNKRIC CHARACTERS.

Cladonemidae with 4 or more simple radial-canals and an equal number <>t biturcated
tentacles. Terminal branches ot tentacles end each in a knob-like. cluster of nemntocysts.
Manubrium a simple, 4 or more sided tube without oral tentacles or prominent lips. Velum
well developed and there is an urticating ridge around the exumbrella side of the hell-margin
below the ring-canal. There is a peculiar brood-pouch above the stomach, but this pouch is
not connected with the gastrovascular cavity of the medusa. The cavity of this brood-pouch is,
however, connected with the bill-cavity by means of simple, interradial openings. The genital
products are developed exclusively in the epithelial lining of this brood-pouch, which is dei i\ i-d

94

MEDUS.E OF THE WORLD.

from the ectoderm of the subumbrella cavity of the bell. The medusa is hermaphroditic; ova
develop in the ventral, and sperm in the dorsal (aboral) wall of the brood-pouch.

The hydroid is Clavatella of Hincks, 1861 (Annals and Mag. Nat. Hist., ser. 3, p. 73,
plates 7, 8; Ibid., 1868, British Hydroids, vol. I, p. 73, plate 12, fig. 2). In Clavatella the
polypites arise singly from a linear stolon. The polypites are small linear, or clavate, with
a zone of 8 slender tentacles which terminate each in a knob of nematocysts. The medusa-buds
are borne only on two opposite sides of the body of the polypite near its base. The hydroid
lives on Ulva in shallow tide pools, and the medusae are produced in summer and autumn.

Hartlaub found that the entoderm of the planula of Eleuthe ria contains numerous nema-
tocyst-cells. Krumbach, 1907, believes that Tnchoplax, F. E. Schulze, 1891 (Abhandl. Akad.
Berlin), is the creeping planula of Eleuthena.

Elfuthcna, Cladonema, and Amphogona are the only genera of hydromedusae known to
be hermaphroditic.

Fit;. 46. — Hydroid (Clavatella prolifera) and its medusa (Eleutheria dichotoma), after Allman, in Ray Society, 1871, 1872.
FIG. 47. — Clavatella prolifera, after Hincks, in British Hydroid Zoophytes.

Eleutheria dichotoma Quatrefages.

Eleulheria dichotoma, QUATREFAGES, 1841, Compt. rend. Acad. Sci., Paris, tome 15, p. 1 68; 1842, Annales Sci.Nat., Ser. 2, p. 270,

plate 18. — KROHN, 1861, Archiv. Naturgesch., Jahrg. 27, p. 157.
Clavatella prolifera (hydroid), HINCKS, 1861, Annals and Mag. Nat. Hist., ser. 3, vol. 7, p. 73, plates 7, 8; 1868, Hist. British

Hydroid Zoophytes, p. 73, plate 12, fig. 2.

Non Eleutheria dichotoma, CLAPAREDE, 1863, Beob. Anat. und Enrwick. wirbelloser Thiere, p. 4, taf. i, fign. 4-10.
Clavatella prolifera (hydroid), ALLMAN, 1872, Monog. Tubul. Hydroids, pp. 31, 212, 384, plate 18.
Eleutheria dichotoma (non claparede), HAECKEL, 1879, Syst. der Medusen, p. 106.
Eleutheria dichotoma, GRAEFFE, 1884, Arbeit, Zool. Inst. Wien, Bd. 5, p. 353. — HARTLAUB, 1886, Zool. Anzeiger, Bd. 9, p. 706,

I fig. Also: Ibid., 1887, Bd. 10, p. 652; 1907, Nordisches Plankton, Nr. 12, p. 127, fign. 1 19, 120. — BEDOT, 1905, Revue

Suisse de Zool., tome 13, p. 133 (literature cited to 1850). — MULLER, 1908, Zeit. fijr wissen. Zool., Bd. 89, pp. 34, 73,

taf. 3, fign. 3-7 (origin and structure of eggs).

This medusa is exceedingly variable, but the normal form may be briefly described as
follows :

Bell irregularly hemispherical, with its lower surface more or less 6-sided. 0.3 to 0.4 mm.
wide. There is a ridge of large nettling warts extending around the margin. There are 6 ten-
tacles at the ends of the 6 short radial-canals, and not irregularly arranged in reference to the
radial-canals as in E. claparedii Hartlaub = E. dichotoma Claparede.

A. \THOM Kill S.K KI.KI TIIKK1A.

95

Each tentacle bifurcates and is twice as long as the bell-diameter. The 2 terminal
branches end one in an adhesive disk, and the other in a large, knob-like cluster of nettle-cells.
These branches are not quite as long as the basal shaft of the tentacle itself. An abaxial
ocellus is found at the base of each tentacle.

The medusa is hermaphroditic, and the sexual products arc developed in an ectodermal
brood-pouch above, but not connected with, the stomach. The cavity ol the brood-pouch is
connected with the bell-cavity by means ot 6 simple, interradial openings which alternate with
the 6 radial-canals. Sperm develops in the aboral, and ova in the oral ( lower) \\all of the
brood-pouch.

Medusa-buds are also produced upon the exumbrella side of the ring-canal, and this
process is usually associated with the sexual reproduction. The terminal suckers of the ten-
tacles are orange or yellowish-brown. Stomach and canals yellowish.

This medusa is found clinging to green sea-weeds oft the Atlantic coasts of Belgium,
England, and France, and is widely distributed in the Mediterranean. Graefte. 1884, found
it to be common upon Ulvu from July to September at Trieste, Adriatic Sea. The budded
medusse become sexually mature in 3 to 4 weeks. The planula larva creeps over the sea-
weeds.

Detailed descriptions are given by Ouatrefages and Allman, and the best modern account
is that of Hartlaub, who discovered the remarkable character of the brood-pouch or ecto-
dermal gonad.

FIG. 48. — Eletithfriti difholiwa, after Hartlaub, in Xordischcs Plankton.

The hydroid is Clavatella prolifera Hincks (see definition of the genus Eleutheria).

Eleutheria dichotoma is apparently quite variable. The tentacles and radial-canals, al-
though normally 6, may range from 4 to 8. Haeckel, 1879, p. 106, designates these aberrations
as "subspecies, " and gives specific names to twelve of them, although a number of these
should he credited to Eleutheria claparcJu Hartlaub and its variations.

Hartlaub finds that the ova develop into planula larvae within the brood-sac before being
set tree.

Miiller, 1908, studied the origin and structure of the eggs of this medusa. The eggs are
small, numerous, and oval in outline, not amcrhoid. There is no visible distinction between
exoplasm and endoplasm. The ooplasma is a network of very fine fibers. There are num-
erous small yolk-granules.

Eleutheria claparedii Hartlaub.

Eleuthrria elafaredii, HARTI.AI-K, 1889, Zonl. Anzeiger, Bil. 12, p. 665; 1907, Noriitscln^ Pl.mkt"ii. Nr. 12, p. 129. fii;. 121.
EIrullifria dichotoma, CLAPAREDF., 1863, Bcobacht. Anat. und Entwicklungsges, \virhrllosrr Thicrr, p. 4. t.if. I, figs. 4-10. —
SPAGNOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 24, tav. 4, fig. 2.

Bell 0.4 to 0.5 mm. wide, irregular in shape, usually more or less hemispherical (fig. 41; i.
A ring of nettling warts on bell-margin. 8 to 10 tentacles, irregularly arranged in reference
to the 4 to 6 radial-canals. A small abaxial ocellus at the base of each tentacle. The ten-
tacles are 3.5 times as long as the bell-diameter, and they bifurcate at their extremities; one
branch ends in an adhesive disk, and the other in a nettling knob. About (> or ~ medusa-buds
are produced upon the subumbrella side of the ring-canal and project into the bell-cavity.

96

OF THE WORLD.

This process of budding; is independent of the sexual reproduction. The terminal suckers ot the
tentacles are orange. Found on Ulva in the Bay of Naples. Described in detail by Hartlaub.
The structure of the reproductive sac is discussed in describing the character of the genus

Eleutheria.

Eleutheria vallentini Brown.

Eleutheria vallentini, BROWNE, 1901, Annals and Mag. Nat. Hist., scr. 7, vol. 9, p. 279.

Umbrella hemispherical, 3 mm. wide, 2 mm. high. 24 tentacles divided into two branches,
the upper branch with clusters of nematocysts, the lower with a terminal sucker. "An ocellus
on the extreme margin ot the umbrella opposite each tentacle." Stomach-tube conical and
small. Mouth a plain round opening without lips. The gonads occupy "the whole of the
upper part of the umbrella above the stomach." A single specimen was found by Vallentin at

Fii;. 49. — Eleutheria claparetlii, after Hartlaub, in Nordisches Plankton.

Stanley Harbor, Falkland Islands, and described by Browne. Color ( ?) Number of radial-
canals ( ?) Brood-pouch ( ?) In the absence of a figure or a more detailed description it will
probably be impossible to redetermine this form unless it be rediscovered in Stanley Harbor.

Genus MNESTRA Krohn, 1853.
Mnestra, KROHN, 1853, Archiv. fiir Naturges., Jahrg. 19, p. 278. — GUNTHER, 1903, Mittheil. Zool. Sta. Neapel., Bd. 16, p. 35.

The type species is Mnestra parasites Krohn, from the Mediterranean.

GENERIC CHARACTERS.

Cladonemidae with 4 radial-canals and a ring-canal. No brood-sac above the stomach.
4 to o degenerate, hollow tentacles with a row of netting capsules along their aboral sides.
\\ ith a ring of nettling cells around the margin and 4 linear tracts of nematocysts over the

ANTIKiMI.IM S.K — MM.STKA.

exumbrella, one above each of the 4 tentacle-bulbs. The throat of the medusa is blocked by
a spongy mass of entoderm. There is a cup-like depression in the center of the exumbrella.

This medusa attaches itself by its suctorial mouth to the throat of the opisthobranch
mollusk, Phyllirhoe. It may lie derived from some Zanclea-like form \\bich has become
degenerate through its sessile habits. It can not swim, yet it has well-developed circular
muscles in the subumbrella and a distinct velum.

Mnestra parasites Krohn.

Mnf*i>, i /Hi'iifitef, KRIUIN, 1X53, Arclnv. Natures., Jalirg. 19, p. 178. — CLAIS, 1875, Vi-rli.indl. /.mil. Buun. Gi-M-ll., \Vu-n.
Bd. 25, p. 9, taf. i. — HAF.C'KEL, 1880, Syst. dor MrduM-n, p. 653. — GUNTHKK, 1903, Mmhnl. Znol. Sta. Ni-a|vl, lid. id.

p. 35, plates l, 3, 42 figs.
Mncstrii parasitica, FEWKES, 1884, American Naturalist, vol. iS, p. 197, figs. 4, 5.

5'

Fid. 50.- — Mnestra parasites, from Florida, after Kc\\kt*s, in Aincr. Natu-
ralist, 1884.

Fn.. 51. — Mnestra parasites^ after Giinthrr, in Mitth. Zool. Sta. Nc.ip
Bell turned inside out, showing iiunit]i-|>.irt -, and cauti
exumbrella to be cup-shaped. Medusa hat. 3 \vcll-»lc\r1f>pi-<l
tentacles.

FIG. 52. — Ctenaria ctenophora, after Hacckc], 1879.

This minute, degenerate, highly variable medusa is parasitic or commensal upon the
mollusk Phvilit'hrjf. Its germ cells appear to migrate into the body of the PAy///rAo« where
they develop.

98 MEDrS.E (IF THE WORLD.

It has been rescued from scientific oblivion through an able paper by Giinther, 1903, who
has carried out an elaborate study of its anatomy, life-history, and variations.

The characters of the medusa are those of the genus of which it is the sole representative
(see genus Mnestra).

Fewkes, 1884, describes this medusa upon Ph\lhrhoc on the Florida Reef. It is com-
mon at Naples and Messina, Mediterranean.

Genus CTENARIA Haeckel, 1879.

Clenaria, HAECKEL, 1X79, Syst. der Medusen, p. 107. — GUNTHER, 1903, Mittheil. Zool. Sta. Nrapel, Bd. 16. p. 57. — HARTLAUB,
1907, Nordisches Plankton, Nr. 12, p. 24.

The type species is Ctenaria ctenophora Haeckel, from the coast of japan. This is the
sole representative of the genus, and Haeckel had but a single preserved specimen.

GENERIC CHARACTERS.

Cladonemidae with 4 bifurcated radial-canals. With 2 feathered marginal tentacles and
simple, unbranched, oral tentacles. There are 8 adradial, meridional lines of nematocysts
over the exumbrella, and a nematocyst tract above the base of each marginal tentacle.
Gonads on the sides of the stomach. There is an apical cavity above the stomach, but it is
not known whether this is an ectodermal brood-pouch, as in Eleutheria, or a mere extension of

the stomach-cavity.

Ctenaria ctenophora Haeckel.

Ctenaria cttnttphora, HAECKEL, 1879, Syst. der Medusen, p. 108, taf. 7, figs. 5-7. — HARTLAI-B, 1907, Nordisches Plankton,
Nr. 12, p. 126, fig. 119.

Bell three-fourths-egg-shaped; 6 mm. high, 5 mm. wide. (Fig. 52.) Widest in a zone
slightly below the middle. 8 adradial lines of nettling cells extend up the sides of the
exumbrella from the margin nearly to the apex, and in addition to these there are 1 large
club-shaped clusters of nematocysts extending from the bases of the 2 tentacles half-way up
the sides of exumbrella.

The tentacles are each 2 to 4 times as long as the bell-height and they taper gradually
from base to tip. There are no basal ocelli. A row of numerous, slender filaments arises from
the abaxial side of each tentacle. These filaments superficially resemble those of Ctenophorae,
and do not end in nematocyst capsules as in Zanclea. They are, however, covered with nema-
tocysts throughout their lengths.

The velum is well developed. 4 main radial-canals arise from the stomach, but these
bifurcate so that 8 canals join the marginal circular vessel in the 8 adradii. The edges of these
radial-canals are serrated, being apparently beset with glands.

The stomach is almost spherical and about half as long as the depth of the bell-cavity.
There are 16 short, simple, oral tentacles, each of which ends in a nematocyst-knob.

Haeckel describes 4 swollen, radially placed gonads on the sides of the stomach, but
these are probably adradial or interradial ( '.) There is a well-developed apical cavity above
the stomach, but it is not quite clear whether this is a brood-sac homologous with that of
Eleutheria or whether it is simply an apical extension of the stomach. Haeckel supports the
latter view, but he made no sections, and the point can not be considered as determined.
Color ( ?)

Haeckel describes this medusa from a single alcoholic specimen from Japan.

Genus CLADONEMA Dujardin, 1843.

Cladoncma, Dl'jARDlN, 1843, Anna!, des Sci. Naturelles, tome 20, p. 370. — KROHN, 1853, Muller's Archiv. fur Anatomie und
Physiologic, p. 420. — GEGENBACR, 1856, Zeit. fur wissen. Zoo]., Bd. 8, p. 230. — VAN BENEDEN, 1866, Fauna Littor.
Belgique, p. 139. — HINCKS, 1868, British Hydroid Zoophytes, p. 62. — ALLMAN, 1871-72, Monograph Tubularian
Hydroids, pp. 216, 357. — HAECKEL, 1879, Syst. der Medusen, p. 109. — PERKINS, 1902, Johns Hopkins University Circu-
lars, No. 155, p. 21;, 1908, Papers from Tortugas Laboratory Carnegie Institution of Washington, vol. I, p. 136. —
HARTLAI-B, 1887, Zool. Anzeiger, Bd. 10, p. 654; 1907, Nordisches Plankton, Nr. 12, p. 131. — GUNTHER, 1903,
Mitt. Zool. Sta. Neapel, Bd. 16, p. ^7.

Stauridium cladontma (hydroid), or PLESSIS, 1888, Recueil Zool. Suisse, tome 4, p. 536.

GENERIC CHARACTERS.

Cladonemidae with 4 or 5 or more bifurcated radial-canals or 8 to 10 or more simple
canals, or with some bifurcated and some simple canals. With 8 to 10 or more tentacles which

AXTHOMKIH S.K- ( I.AIHINI.MA. (I'.l

give rise to sucker-bearing or nematocyst-bearing branches or both. With 4 or 5 simple, oral
tentacles which terminate in nematocyst-knobs. There is no brood-sac above the stomach.
The genital products develop within the entoderm of the stomach, and when mature are
found in the ectoderm. The medusa is sometimes observed to be hermaphroditic.

The type species is ClaJonctna radiatum of the coasts of Europe. According to Haeckel
this species is highly variable both in form and color (see Haeckel, 1879, p. 109). The two
American forms described by Perkins are, however, quite constant in their form, although
one of them is somewhat variable in color. The hydroid stock of ('.laJuni-nm belongs to the
tubularian genus StauriJia of Dujardin. It is well described by Allman, 1871 (Tubularian
Hydroids, pp. 216, 357, plate xvn), and by Perkins, 1908.

Cladonema radiatum Dujardin.

Cladonema radiatum, Dt'jARDiN, 1843, Compt. rend. Acad. Sci., p. 1134.

Siauridie (hydroiil), DUJARDIN, Ibid., p. 1133.

C.ladonema radiatum and Stauridie, DIIJARIMN, 1X43, Annal. des Sci. Nat., tome 20, p. 370; 1845, Ibid., ser. 2, tome 4. pp. 271
272, plate 14, fig. C.

C.ladontma radiatum, KROHN, 1853, Muller's Archiv. fur Anat. und Physiol., p. 420, plate 13. — KKFKRSTFIN i ND EHI. F, RS, 1861 .
Zoologische Beitrage Neapel, Messina, p. 85, taf. 13, fig. 5.— VAN BENEDEN, 1866, Mem. Acad. Roy. Belgnjue, tome 36,
p. i 39, plate 12. — HINCKS, 1868, Hist. British Hydroid Zooph., p. 62, plate 1 1 . — ALLMAN, 1872, Monog. Tubul. Hydroids,
pp. 216, 357, plate 17, figs. l-io. — HAECKEL, 1879, Syst. der Medusen, p. 109. — WEI s MANN, 1883, Entstehung Sexual/ ellen,
bei Hydromedusen, Jena, pp. 1 19, 218, taf. 12, figs. 2-5. — JICKELI, 1883, Morphol. Jahrbuch, Bd. 8, p. 602, taf. 26 (his-
tology).— HARTLAIIB, 1887, Zool. Anzeiger, Bd. 10, p. 655, I fig.; 1907, Nordisches Plankton, Nr. 12, p. 132, fign. 123-125,
(list of authors; excellent description of the medusa). — BEDOT, 1905, Revue Suisse de Zool., tome 13, pp. 58, 132 (refer-
ences to 1850). — BILLARD, 1905, Bull. Museum d'Hist. Nat. Pans, tome ll, p. 500 (variations). --Mi'LLi R, 1908, Zrit.
fur wissen. Zool., Bd. 89, pp. 30, 73, taf. 3, figs, i, 2; taf. 4, figs. 8-1 1 .

Bell half-egg-shaped or globular and 2 to 3 mm. wide. At Naples, Italy, the medusa
appears to be quite invariable, and according to Haitlaub, 1887, the manubnuin is spindle-
shaped and five-sided with 5 perradial oral tentacles and 5 perradial sac-like outgrowths
upon the sides of the stomach. Thus in the Naples medusae 5 main radial-canals, 72° apart,
arise from the stomach, but 3 of these bifurcate so that 8 canals, 45° apart, join the circular
vessel at the margin. These 5 main radial-canals are arranged as follows: 2 bifurcated
canals 72° apart; 2 simple, unbranched canals 144° apart; i bifurcated radial-canal mid-
way between the two simple canals, i. e., 72° from each. Thus in glancing around the
margin in the direction of the order of succession of the hour-numbers on the dial of a clock,
we may begin with a simple canal, then comes a bifurcated, then a simple, next a bifurcated,
and finally another bifurcated canal. 2 of the bifurcated canals are thus side by side and
the 2 simple canals are separated one from another by an interval occupied by the third
bifurcated canal.

There are 8 tentacles upon the Naples medusae, one at the end of each terminal radial-
canal. The basal bulbs of these tentacles are heavy and swollen, and each bears an abaxial
ocellus, i to 3 sucker-bearing filaments arise from the inner sides of the tentacles and their
outer parts terminate in 4 to 6 branches which are armed with nettling warts and end in
knobs.

The gonad encircles the stomach and is not confined to the sac-like protrusions, as was
believed to be the case by Haeckel. According to Hartlaub, the genital products originate
in the entoderm, but when mature are found in the ectoderm of the manubrium; yet according
to Weismann they originate and remain in the ectoderm.

The medusa exhibits a successive hermaphroditism, according to Hartlaub, although
either sex may precede in the process. (See also Miiller, 1908.)

The stomach, ring-canal, and tentacles are red to brown.

This medusa is abundant oft" the Atlantic coasts of England, Holland, and V ranee, and
in the Mediterranean. Haeckel records a number of departures from the normal form.
These are probably only aberrations, but he gives specific names to each and every one of
them. For example, the oral tentacles and the sac-like outgrowths on the stomach may range
from 4 to 5; and there may be 10 simple, or 4 bifurcated radial-canals.

There appear indeed to be many local races of this medusa, and I am inclined to believe
that Cladonema perkinsii and (<. nmyen of America may prove to be only varieties of C.
radintuin.

100

MK.WS.E OF THK WORLD.

Billard has studied the variations of the medusa at the Bay de la Hougue near St. Vaast
on the northwestern coast of Fiance. Here he found that among 50 medusae 5 had 6 radial-
canals, 4 had 7, 39 had 8, I had 9, o had 10, I had u. In the case of the 39 medusae with
8 radial-canals, 36 had 2 simple and 3 bifurcated canals, as in the Naples medusa.

Miillei, 1908, has studied the origin and structure of the ova in this medusa. The eggs
are small and rounded and widely scattered in the gonad, for the successful eggs devour
the weaker in the ovary. The ooplasma is sparsely vacuolated and there is very little yolk.
53- 55-

54-

";; .- Hydroid of ClaJonema raiiiatum, after Hincks, in British Hydroid Zoophytes.
Showing growth-habit of hydroid and enlarged view of single polypite.
Fir.. 54. — ClaJonema raJiatum, after Allman, in Ray Society, 1871-72. Budding

hydroid anil mature me lusa.
Fi<;. 55. — Medusa of C.lailontma rnJiatum, after Hincks, in British Hydroid Zoophytes.

The hydroid of Clndoncinti radiattitn isStaurn/ia raJiatum Dujardin = CAndonema radiatum
Hincks.

The hydrorhiza is a slender, creeping filament from which the hydranths arise at inter-
vals. These hydranths are either simple, unbranched, and about 2.5 mm. high, or are borne
upon slightly branching stems 12 to 25 mm. high. The hydranths are club-shaped and
have 2 alternating verticils, each circlet consisting of 4 tentacles. The circlet near the mouth
is composed of long tentacles, each of which ends in a knob, while the circlet at the base of
the hvdranth consists of 4 very short, simple, unbranched, filiform tentacles which alternate
with (i. e. are upon meridians 45° apart from) the oral circlet. The medusae bud out singly
from the sides of the polypite at a short distance above the basal circlet of tentacles. The

\\Tim\ii.Di s i; CI..\I>H\I:M \. 101

body of" the hydranth Is pale reddish and the perisarc bright yellowish-brown. The medusae
are produced in spring and summer.

Dujardin's, 1843, observations upon Cladonema niJintiun are the earlit-st in which the
complete life-history ot' the alternation of generations between hydroid and medusa was
actually observed.

Cladonema perkinsii Mayer.
Plate 9, fig. i .

C.lailonema, sp., PERKINS, 1902, Johns Hopkins Uimersitv Circulars, vol. n. No. 1^5. p. 2^, ne,^. in ti-it.

Cladonema per kin sii, MAVFR, 1904, Memoir- N.it. Sci. Brooklyn Inst. Arts ami SM .. vol. i , XL. i . p. i X, plate 4, fig. 35. — H*KT-
LAi'B, 1907, Nonlisches Plankton, Nr. 12, p. i ;^, tit:, i±t>.

Bell half-egg-shaped; less than 2 mm. in diameter, with thin, uniform walls. 8 lar<;e,
stout, marginal tentacles. The inner and lateral parts of each of these tentacles bear 3 to 10
small, flexible cirri, which are besprinkled with wait-like clusters of nematocysts and termi-
nate in a knob. 1 here is a large, cup-like, ectodermal ocellus upon the outer side of each
main tentacle shaft near the bell-margin. The velum is large and shows circular striations.
Manubrium large and spindle-shaped, with about n rounded, protruding pouches at its widest
part. The mouth is surrounded by a circlet of 5 simple, short oral tentacles each terminating
in a knob-like mass of nematocysts. 8 simple radial-canals, 45° apart, arise hum the stomach,
and extend straight unvard the simple circular vessel. The manuhrium is thus 8-sided at
its proximal end, 6-sided in the middle, and 5-sided at the mouth. The genital products
develop in the ectoderm ot a large part ot the manubriurn and also in the hernia-like pouches.
The ocelli are black and all other parts colorless. This species was discovered by Professor
Perkins in Nassau Harbor, Bahamas, in July, 1902, upon the surface in shallow water at
night.

Cladonema mayeri Perkins.
Plate 9, figs. 2 an, I -, .

Cladonema, .</>., FKWKES, iSS^, Bull. Mus. Comp. Zool. at Harvard College, vol. 11, p. 87.

Cladonema ma\eri, PKRKINS, 1906, Year Book of tile Carnegie Institution of Washington, No. 4, IIJCK. p. llS. — KjoS, Papers
from Tort u gas I. ah. Carnegie In ^t. Washington, \ol. i , p. i ;(>, plate- i an, I 2. plate 4, figs. 2 t , 22. H\ ,!roul an, I medusa.

Bell thin-walled, higher than a hemisphere, and with small, solid, apical projection;
top rounded and dome-like, and total height about 2.5 mm. There are 9 marginal tentacles,
one at the toot of each radial-canal. Each ot these tentacles has a large, spindle-shaped basal
bulb, the entoderm of which contains a mass of white concretions. On the outer side of
each basal bulb near the circular canal there is a deep reddish-brown, ectodermal pigment
spot. The distal, inner side of each basal bulb gives rise to about 6 small tapering peduncles
which terminate in small knob-like adhesive disks. These enable the medusa to cling to the
sides or bottom of the aquarium. The main shaft of each tentacle extends outward from the
spindle-shaped end of the basal bulb. It is umtorm, thread-like, and slender, and gives rise
to 4 to 8 thread-like side branches each ot which, together with the central shaft, terminates
in a knob-like cluster ot nematocvsts. There are 4 to 6 nematocyst-warts upon each side
branch and a greater number upon the main shaft. Both the main shaft and the side brandies
are highly contractile and can be expanded so as to become longer than the bell-height, or
contracted into a close bunch. The velum is very wide and its orifice small. 6 radial-canals
arise from the stomach, but every alternate canal bifurcates near its point ot origin, and
thus () equally spaced canals reach the circular vessel. The manubrium normal!) extends
to the level of the velar opening, but may contract somewhat at times, as is shown in our
figure. The mouth is surrounded by <> simple oral tentacles, each of which terminates in
a large nematocyst-knob. Near the middle of the stomach there is a circlet of 6 radially
arranged, short, blunt, hernia-like projections. The genital products are developed in the
ectoderm of the walls of the manubrium.

The color is quite variable. The entoderm ot the basal bulbs ot tin- marginal tentacles
is milky-yellow streaked with dark-brown pigment-granules, while the entoderm of the
manubrium is milky-yellow or dull-milky ocher, streaked longitudinally in each of the ft
radii by dark-brown, almost black, pigment-granules. The disposition and arrangement ot
this pigment is highly variable. Perkins, iijoX, nives the most complete account ot this
medusa and its hydroid.

102 MF.nrs.K UK TIIK WORLD.

This medusa, together with its hydroid stage, was found in great numbers by Prof.
Henry F. Perkins in the salt-water moat of Fort Jefferson, Tortugas, during the summer of
1905. The hydroid is a minute Stauridiu which grows upon algae. The same medusa was
described by Fewkes, 1883, as being in association with Cassiopea at Fleming's Key, near
Key West, Florida.

Genus DENDRONEMA Haeckel, 1879.

Dtmlronema, HAFCKEL, 1879, Syst. der Metlusen, p. MO. — GUNTHER, 1903, Mitth. Zool. Sta. Neapel, Bil. 16, p. 57.

The only known form is Dendronema styludendron Haeckel, from the Canary Islands,
Atlantic Ocean.

GENERIC CHARACTERS.

Cladonemidae with branched oral tentacles and branched marginal tentacles, the
branches ending in nematocyst-knobs or adhesive disks, or both. With bifurcated radial-
canals. Gonads in the stomach-wall.

There is an apical cavity above the stomach, but we do not know whether this is a mere
extension of the stomach itself, or a reproductive sac similar in anatomy to that of Eleutkena.

Dendronema stylodendron Haeckel.

])?rnlionetiiti st \lrjiletidT on, HAVCKKL, 1879, Syst. iler Medusen, p. no, taf. 7, fi^. 8.

Bell miter-shaped, with pointed conical apex. 9 mm. high, 6 mm. wide. Stomach spindle-
shaped. There are apparently 4 radially placed oral tentacles each of which branches dichot-
omously 6 to 7 times and terminates in (50 to 60) nematocyst-knobs in each quadrant.
Haeckel's description is, however, vague upon this point. Haeckel states that there are 4 egg-
shaped "gonads" on the 4 perradial sides of the stomach. These may, however, be homol-
ogous with the protuberances seen in Cladonema ( ?) HaeckePs figure shows them distended
with ova. 4 principal radial-canals, 90° apart, leave the stomach and bifurcate so that 8
vessels reach the ring-canal 45° apart. There are 8 marginal tentacles, one at the base of each
radial-canal. There is an ectodermal abaxial ocellus at the base of each tentacle. Each ten-
tacle gives off a stout, short branch on its inner (velar) side, .and this branch bifurcates twice
and terminates in 4 nematocyst-bearing, knob-like "suckers," or adhesive organs. The main
shaft of the tentacle also branches dichotomously many times and is besprinkled with nettling
warts, and the ends terminate in large knobs. There is a spindle-shaped cavity in the apex
of the bell, above the stomach, but its anatomical character is unknown. The mouth, stomach,
apical cavity, and gonads are reddish-yellow. Tentacles, radial-canals, and ring-canal brown-
ish-red. Ocelli black.

This medusa was found by Haeckel in the Canary Islands, Atlantic Ocean, in February,
1867.

Family OCEANIDjE, sens. Vanhoffen, 1891.

OeeaniiLr (in part), ESCHSCHOLTZ, 1829, Syst. der Acalephcn, p. 96. — FORBES, 1848, British Naked-eyed Medusa-, p. 21.—

GEGENBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 219.

Tiarid.e+ Margt'lidlp + Cannotidtf (in part), HAECKEL, 1879, Syst. der Medusen, pp. 40, 68, 1^6-159.
Ocrnniiir, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 443.

FAMILY CHARACTERS.

Anthomedusa? with 4, or 4 pairs of, isolated gonads upon the interradial or adradial sides
of the stomach. The marginal tentacles may arise singly or in clusters, but are neither
branched nor feathered.

The Oceanidae constitute the third family of the Anthomedusae, and they are more com
plex and more highly differentiated than the simple and more primitive Codonidae.

AXTHOMKlll S.K -OCEANID.S. HIM

We distinguish three subfamilies ot" Oceanidae.

1. Tiarirue. Lips without oral tentacles. Simple unbranched radial-canals. Ten-

tacles arise separately from the hell-margin and are not grouped in clusters.

2. Margelinti'. With oral tentacles. Simple, unhranched radial-canals. In some

genera the marginal tentacles arise singly; in others they are grouped in clusters.

3. DenJrostaurtnte. Lips without oral tentacles. The radial-canals branch.

Direct development ot the medusa trom the egg is unknown in this tamilv. The following
genera are known to develop through Tubularian hydroids: Stomotoca, Turrn. Podocaryne,
Stylactis, Bougainvillia, Neinopsts, Lizzia, and ftillia.

Medusae are produced asexually upon the interradial sides of the manubrium in ('\tttis,
PoJocoryne, Bougainvillia, and Rathkea, In some species of Bougainvillia the eggs undergo
part of their development within the ectoderm of the parent medusa, and are discharged as
well-developed planulae. In Ntobia the tentacle-bulbs develop into new medusae, and are set
free one after another to repeat this process. In H illia and Prohoscidactyla, medusa-huds
are produced upon hollow stolons which may arise from the corners ot the stomach at the
points ot |uncture with the radial-canals, or from the places where the canals fork, as in P.
flavicirrata, var. stolonifera, Maas.

Haeckel, 1879, describes the gonads ot the Oceanidae as being penadial in position anil on
the sides of the stomach in the same sectors with the radial-canals, but Vanhoffen, 1891 (Zool.
Anzeiger, Bd. 14), showed that this is an error, for the gonads are almost universally adradial
or interradial in position and alternate with the sectors of the radial-canals.

The specialized conditions displayed by the medusae ot the Oceanidae, such as the corru-
gated, folded gonads ot the Tiannae, the clustered marginal tentacles and the oral tentacles of
the Margelmae, and the forked radial-canals and other peculiar structures in the Dendro-
staurinae.all indicate that the Oceanidae are derived from simpler forms, such as the Codonidae.
Indeed, the conditions seen in theOceanidae are largelytoreshadowed in the more differentiated
Codonidae and in the Cladonemidae. In common with the Codonidae and Cladonemidae
the Oceanidae are derived trom Tubularian hydroids.

Subfamily TIARINjE Haeckel, 1879.

(liemnd.r (in part), ESCHSC HOLTZ, 1829, Syst. tier Acalephen, p. <)(>. GK^NBACR, lNcf>, 7.ni. fiir uissrn. /mil., Hd. S,p. 219.

\mleiffrtF (in part), LKSSON, 1X45, Hist. Nat. Zooph. Acal., p. 28}.

Tjitridii-, HAF-CKRL, 1879, Syst. der Medusen, p. 40.

Sfns. reslr,, drlomrrinthiu, VANHO? UK, 1891, Xool. Anzeiger, Bd. 14, p. 44V

Tiiiriiiit, HARTLAUB, 1892, Nachrichten kgl. Gesell. Wissenschafl. Univ. Gottingen, pp. 19-22; 1907, Nordisihes 1'l.inkton,

Nr. 12, p. 5.

.SVfn. restr.t Tiurijcp, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. iS, |>. ri.
Tiilrin.r, VON LENDENPELDj |NS4, X.uol. An/fiytT, Bd. 7, p. 446.

CHARACTERS OK THF. TIARIN/E.

Anthomedusae with 4 lips, without oral tentacles. With 4 or more unbranched radial-
canals and with simple, hollow, unhranched tentacles which are not grouped in clusters, but
arise separately from the bell-margin. With interradial or adradial gonads forming swollen,
corrugated regions in the walls of the stomach.

The following table shows the genera of the Tiarinae.

1 . Tnhc PROTIARIDI :

The outer surfaces of the four interradial gonads are smooth. Not corrugated or folded. There are 4 simple lips, 4 r.uii.d-
canals, and a ring-canal, 4 or more tentacles.

Proriara HAtcKEL, 1879. With 4 radial-canals and 4 radially situated marginal tentacles. 4 simple lips not com-
plexly folded. Hydroid unknown.

Hetrroliara MAAS, 1905. With 4 radial-canals, and 8 marginal tentacles. Ring-canal gives rise to interradial, blindly-
ending, centripetal canals. Only species is H. anon\mn, MAAS, 1905, Craspedoten Medusen Siboga Expedition,
Monog. lo, p. 19, taf. 3, figs. 19-21, Malay Archipelago. Hydroid unknown.

2. Tribe AMPHINEMIDI:

With 2 long and numerous rudimentary tentacles. There are 4 radial-canals.

Slomoloca L. AOASSIZ, 1862 = Amphincma + Stomoloca+ Codonorchis, HAICKII, 1^79. With two well-developed ten-
tacles. Manubrium mav or may not be mounted upon a peduncle. Gonads corrugated, folded, swollen regions upon
interradial or adradial sides of stomach.

Dissontma HAECKEL, 1879. With 2 well-developed and numerous rudimentary tentacles. During their development
gonads migrate outward from sides of stomach down the 4 radial-canals.

104 MKDT'S.K OF THE WOULD.

V Tribe PAND.IIDI:

With 4 or more well-developed tentacles. The gonails are interradial, corrugated, or folded ridges in the wall of the stomach

4 radial-canals.
Pundta LESSON, 1843 (1837?). 4 interradial gonads in stomach-wall, not completely separated in the 4 principal

radii. Hydntid unknown.
Conis BRANDT, 1838. Ocelli borne upon ends of special, short clubs which arise from bases of tentacles. Hydroid

unknown.

Turris LESSON, i^^=Tiara, LESSON -4- Catablema, HAECKEL. With 4 interradial crescent-shaped gonads in the ecto-
derm of the stomach-wall. Each crescent is composed of partially fused, swollen ridges. Hydroid: Clavttla ( ?),
Carnpaniclava ; or Perigonimus ( r)

4. Tribe CAI.YCOPSIDI:

With more than 4 simple, unbranched radial-canals. Adradial, transversely folded gonads.

Cnhcopsis FEWKES, 1882. With 16 simple, separate, unbranched radial-canals, 4 radial, 4 interradial, 8 adradial.

Haeckel, 1879, established the family Tiandae for Anthomedusa? with 4 wide, crenulated
lips; with 4 separated or 8 cleft gonads in the stomach-wall; with 4 wide, hand-like radial-
canals; and with simple, unbranched tentacles.

Vanhoffen, 1891, showed that the gonads were interradial, not radial in position as was
supposed to be the case by Haeckel; and in 1892 Hartlaub gave important details of the
structure of the gonads, showing that Panden had simpler gonads than either Tiara or Turris.
For example, he showed that the gonads of Pandea conica consist of 4 interradial, horseshoe-
shaped, network-like, swollen regions in the ectoderm of the stomach-wall. The gonads of
T urns cceca, however, consist not only in the interradial network, hut chiefly in a double row
of fused longitudinal swellings in each interradial quadrant of the stomach. In Tiara pileata,
on the other hand, we find none of these network-like gonads, but instead a horseshoe-shaped
gonad in each interradial quadrant, the apex being upward and the sides of the horseshoe
being made up of a series of laterally-fused, horizontal swellings. Maas, 1904, loc. cit., gives
clear figures of these conditions and supports Hartlaub's observations.

In the more complex and specialized Tiarinae, represented by the tribes Amphinemidi and
Pandaeidi, the gonads are thrown into complex folds or corrugations, and the lips have become
recurved, with folded edges.

Vanhoffen concluded that Amplnnema and Codonorchls of Haeckel are equivalent to
Stomotoca L. Agassiz; and in this I concur. He also maintained that Pandea Lesson was
identical with Tiara Lesson, but in 1892 Hartlaub showed that the gonads of Pandea were
quite different in structure from those of Tiara; and Maas, 1904, supports this conclusion.

Maas, 1904, calls attention to the fact that there are no definite distinctions between
Turns and Tiara. He shows that the gonads of Turris consist of 4 interradial horseshoes, the
sides of each horseshoe being made up of partially fused, transverse, branched, ectodermal
ridges and the upper apex of the horseshoe being composed of an open network of ridges. In
Tiara the gonads are horseshoe-shaped, but commonly lack the network-like ridges. The
transverse ridges do anastomose to some degree, however, in Tiara, so that a partial network is
often found. See Maas, 1904, lor. at., plate 2, fig. II.

Maas describes the gonads of Catablema Haeckel as horseshoe-shaped and composed of
partially fused, vertical ridges. In the fully-grown medusa, however, I find that the ridges
tend to become transverse, as in Turris or Tiara.

Maas, 1904, retains the genus Catablema of Haeckel to include medusae resembling
Turns or Tiara, but with blindly-ending, lateral diverticula upon their radial-canal and ring-
canal. As a matter of fact these diverticula are highly variable in different individuals of the
same species and are seen in a more or less well-developed condition in the majority of Tiarinae.
I therefore believe that confusion will be avoided it we combine the genera Turris, Tiara, and
Catablema to form a single genus. This should be called "Turris," for Lesson used this name
on page 283 of his Hist. Zooph. Acal., and on page 17 of his "Prodrome," 1837 Tiara he
defines later on page 286 of his "Histoire," 1843, and on page 20 of his " Prodrome, "1837.

I propose, therefore, following the lead of Vanhoffen, Hartlaub, and Maas, to reduce the
13 genera of Tiarinae enumerated by Haeckel, 1879, to 8, as follows: Stomotoca, Modeeria,
Protiara, Heterotiara, Pandea, Turns, Corns, and Calycopsis.

The Tiarinae are widely distributed, but are abundant only along continental coasts, for
in so far as is known, they arise by budding from Tubularian hydroids of the genera Pcngoni-
nius, Clavula ( ?), and Campaniclava ( r). Asexual budding or direct development in the
medusa-stage is unknown.

ANTIKiMI Dl - E -I'lioTIAUA. ] Of)

The medusae <>t the various genera of Tiarinae hear a close resemhlance one to another.
Their hells are usually miter-shaped; ectodermal ocelli are often found upon their hollow ten-
tacle-bulbs, and their radial-canals arc usually hroad and Hat, and often with more or less
jagged outlines.

All of" the tentacles arise from the lower edge of" the bell-margin when young, but as
growth proceeds, the upper parts of" the basal bulbs of the older ones are crowded and forced a
short distance up the sides of the bell, while the smaller tentacles still remain upon the lower
edge of the hell-margin. 1 his gives the appearance of two rows of tentacles.

Hartlaub demonstrated that the so-called mesenteries of I lacckel. 1X70,, arc only the wide.
funnel-like origins of the radial-canals, where they communicate with the stomach-cavity.

As Maas, 1904, has shown, the Tiarinae have given rise to the more specialized Bytho-
tiandi, wherein the radial-canals have become branched, and the latter arc probably related
to the \\illiadi. I he Iiarmas are themselves derived, probably, from Codonidae in which
the originally ring-like goiuul has become radially separated, so that it lies only in intrn.uli.il
positions on the wall of the stomach. They are thus, apparently, more highly specialized
than the Codonidae. Cnl\; »f>m with its 16 simple, unbranched radial-canals may be regarded
as a form intermediate between the Tiarinae and Bythotiaridi.

The Tiarinae are distinguished from the Margehnae by the tact that oral tentacles are
never found in the Tiarinae, but are present in the Margelm;e. Moreover, the tentacles of
the Tiarinae arise singly from the bell-margin, and are not grouped in clusters as is frequently
the case in the more specialized Margelinae. It seems not improbable that the Tiarinae and
Margelinae have arisen independently of each other from the Codonidae. A decided difference
between the Tiarinae and Margelmae is that in the former the entodermal cores of the tentacles
are hollow, and in the Margelinae they are nearly, it" not wholly, solid. When present the
ectodermal ocelli in the Tiarinae are on the untt'r sides of tin- tentacle-bulbs, whereas in the
Margelinae they are on the inner (velar) sides

Genus PROTIARA Haeckel, 1879.

Carminrolhe btroe, SLABBER, 177?, Physikal. BHust., p. 64.

Proliara, HAECKEL, 1879, Svst. ilrr MciluM-n, p. 46. HARC.ITT, 1902, Biological Bulletin, Boston, vol. 4, p. 17; )<)O4, Bull.

I'. S. Bureau of Fisheries, vol. 24, p. 34. — LINKO, 190:, /.»< -1 . Xii/rim-r, J.ilirt,-. : ^, p. i (12.
( ? ) PlolorniJe, WAI, NCR, 18X5, \VirMlosrn des WIT.M-H Mo-res, BJ. i, p. 74.
Halititirti, KFWKFS, 1882, Bull. Museum Conip. /mil. at Harvanl Collet;.-, M.|. >), p. J-M. V.\^i/ .in.) M^YJR, lS<|i), Hull.

Museum Conip. Zool. at Harvard College, vol. 32, p. 160.

GENERIC CHARACTERS.

Tiarinae with 4 well-developed, radially situated tentacles with hollow, basal bulbs. \\ ith
4 longitudinal, swollen gonads on the 4 interradial sides fit the stomach. The outer surfaces
of these gonads are smooth, not folded, nor corrugated, 1 he 4 lips are simple, not folded noi
crenulated.

Haeckel, 1^79, founded this genus for P. t<ir<i/i<-ni<i, which hail been previously described
by Slabber, 1775, under the name Carminrothe /vro, from the North Sea and Knglish Channel.
According to Vanhorfen, itfiji (X,ool. Anzeiger, Bd. 14, p. 441), this medusa is only a young
('rjr\'iiitis. However this may be, Hargitt. K;oa, discovered a medusa in Yimvaid Sound,
Massachusetts, which accords well with Haeckel's definition of Protiara. The gonads are
described by Hargitt as being found in tour separate, longitudinal, swollen regions in the
interradial (radial ?) sides of the stomach. Hargitt cut no sections of the medusa, and con-
sequently we must merely place this species provisionally in the genus Protiara, for it the
gonads be developed so as to completely surround the stomach and are not separated radially,
the medusa is one of the Codonidae. It it be one of the Tiarinae, i{ appeals to constitute
an interesting intermediate form between some Corynitis like member of the family Codonidae
and the Tiarinae. It has the simple mouth, narrow canals, and smoothly rounded external
surfaces of the manubrium characteristic of the Codonidae, but its 4 separated, interradial
gonads, and its hollow, tapering tentacles allv it to the Tiarinae.

Lmko, IQO2, sectioned a somewhat similar medusa from the Murman coast, between
Russia and Norway, and demonstrated that the 4 gonads are interradial.

The medusa described by Fewkes as f fulitnun fnrninsa is evidently a I'lotiiim.

100 MKDl'S.E OF THIO WORLD.

Protiara^beroe.

Carminrothe beroe, SLABBER, 1775, Physikalische Belustingungen, Niirnberg, p. 64, taf. 14, fig. I.

Oceania tetranema, PERON F.T LESUEUR, 1809, Ann. du Mus. Hist. Nat., tome 14, p. 347. Paris.

Protiara tetranema, HAECKEL, 1879, Syst. der Medusen, p. 47.

(?) Protiara, LINKO, 1902, Zool. Anzeiger, Bd. 25, p. 162, 2 figs.

( >)Proliara borea/is = Phtocnide borealis, Colorless variety, WAGNER, 1885, Wirbellosen des Weissen Meeres, Bd. I, p. 74, taf. 4,

fign. I, 2.
( ?) (Svndictian ?) incerturn, LINKO, 1900, Travaui Soc.Imp.Nat.de St. Petersbourg, tome 29, p. 151, fig. l (this is possibly

Sarsia flannnea ?).
( fyPlotocnidc incena, HARTLAUB, Nordisches Plankton, Nr. 12, p. 70, fig. 66.

Bell 6 to 15 mm. high, 4 to 12 mm. wide. Egg-shaped, with very thick walls, which in
the upper part of the bell are one-third to one-fourth as thick as the hell-height. There are a
few scattered nettle-cells over the exumbrella, these being more numerous in young than in
old specimens. 4 tentacles, each 4 to 5 times as long as the bell-height, with thick basal
bulbs about one-sixth as wide as the greatest width of the bell. No ocelli. The shafts of the
tentacles bear garland-like pads of nettle-cells. There are 4 straight, narrow radial-canals
with smooth edges. The manubrium is short and even in mature medusae does not project
beyond the velar opening. In young medusas it is conical, in mature individuals very wide,
but it narrows greatly at the neck immediately above the mouth. The neck is tubular, and
the mouth is encircled with nematocysts. There are 4 longitudinal, interradial, ectodermal
gonads, which project widely outward in the upper portion of the manubrium, but do not
extend to the mouth. The outer surfaces of the gonads are smooth.

Linko, 1902, sectioned the manubrium and found that the entoderm forms 4 interradial
partial septa which project inward in 4 longitudinal folds toward the axial center of the
stomach, but their inner edges do not fuse. The cells of these septa are digestive and they
serve apparently to increase the area of the stomach-wall. The bell is colorless. Manu-
brium and tentacle-bulbs orange to yellow, radial-canals white.

Common in Barents Sea, North of Russia, between 68° 54' and 70° 5' N. lat.; and
33° 3°' to 57° 38' l°ng- E- from Greenwich. Rare in harbors and fjords.

(Syndiction?) incerturn, Linko, 1900, may possibly be identical with P. beroe. It has
a bell 3 mm. high and somewhat more than 3 mm. wide. The walls are thick and there is a
rounded, dome-like apex sharply set off from the bell itself. The exumbrella is besprinkled
with quite regularly and widely spaced nematocysts. There are 4 thick, tapering tentacles
about one-third to one-half as long as the bell-height. These have large, spherical, non-
ocellated, basal bulbs. Velum well-developed. 4 straight, slender radial-canals. Stomach
mounted upon a short, conical peduncle. The mouth does not reach the level of the velar
opening. Mouth-opening round, without prominent lips. The gonad is figured as being much
swollen above, tapering below, and encircling the stomach. It is not wholly clear from Linko's
description whether there are 4 interradial gonads or only one encircling gonad, for he states
that the medusa resembles P. borealis = (Syndictton boreale Birula) in some of its characters.
Only about 10 very large eggs are produced by the female.

The manubrium is yellow to orange, tentacle-bulbs red, and tentacles yellow. Other
parts are colorless. Found in the White Sea, Northern Russia. It is probable that this form
is a Sarsia, and is possibly S. flammea. See Hartlaub, 1903, 1907. There is, however, no
peduncle in .V. flammea.

Protiara borealis.

Phtocnide borealis, WAGNER, 1885, Wirbellosen des Weissen Meeres, Bd. I, p. 74, taf. 4, fign. I, 2.

Synilictyon boreale, BIRULA, 1896, Annuaire Musee Zool. Acad. Imp. Soc. St. Petersbourg, tome I, p. 336 (Russian).

( ?) Protiara h<fckelii, HARGITT, 1902, Biological Bulletin, vol. 4, p. 16, fig. 4; 1904, Bull. I'. S. Bureau of Fisheries, vol. 24,

p. 34, I fig.
Phtocnide borealis, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 69, fig. 65.

Bell is about 3 mm. high and 1.5 mm. wide, with simple rounded apex and thick walls.
4 radially situated tentacles, each about as long as the bell-height. These tentacles are
besprinkled with nematocysts and have well-developed, conical, basal bulbs, without ocelli.
Wagner's specimens were apparently much younger than Hargitt's and each tentacle termi-
nated in a large knob-like cluster of nematocysts. In Hargitt's specimen the tentacles taper
gradually to their tips. The velum is well-developed. There are 4 straight, narrow radial-

AXTHOMEDL'S-E — PROTIARA, HKTKROTIAHA.

107

canals, and a simple ring-canal. The manubrium is mounted upon a short, conical peduncle
and is quite wide; rectangular above, hut nearly circular in cross-section near the mouth.
There are 4 very small, simple lips. In Wagner's specimens the manubrium was only about
halt as long as the depth of the bell-cavity, whereas in Hargitt's the
mouth extended nearly to the level of the velar opening. This differ-
ence may be due to age(?) The gonads are 4 large, interradial,
longitudinal swellings on the sides of the stomach. The outer sur-
faces of the gonads are smooth.

Wagner found a few specimens of this medusa in April, in the
\\ hue Sea, north coast of Russia. Hargitt found an apparently iden-
tical medusa in summer near No Man's Land, an island off the
southern coast of Massachusetts. The gonads and tentacle-bulbs in
Hargitt's medusa are milky in color, other parts being transparent.
This species is distinguished from Protiara bcroe ( = P. " trtrn-
iii-iiin" Haeckel) by its absence of color and lack of ocelli.

Unfortunately no sections have been made of the manubrium.
Wagner's figure and description are unsatisfactory in that he leaves
kaiekeli," tne position of the gonads in uncertainty and we can not tell whether
there are 4 (interradial ?) gonads or only I encircling gonad.

FK;. 56. — Protiara

after Hargitt, in Biologi-
cal Bulletin.

Protiara formosa.

Plate 6, figs. 4, 5, and 6; plate 13, figs, l and 2.

Halitiara fortnosa, FEWKKS, 1882, Bull. Mus. Com p. Zool. at Harvard Coll., vol. 9, p. 276, platr 4, fig. 2. -MAYK.R, 1904, Memoirs
Nat. Sci. Mus. Brooklyn Inst. Arts and Sci., vol. I, No. I, p. 8, plate I, fig. 8. — A(IASSIZ, A., AND MAYER, 1899, Bull. Mus.
Comp. Zool. at Harvard Coll., vol. 32, p. 160. — MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 37, p. 31.

Bell about 3 mm. high and pear-shaped with solid apical projection. There are 4 long,
tapering, radially situated tentacles, about two-thirds as long as the bell-height. These ten-
tacles are hollow, with long, tapering, basal bulbs, and their ends are usually coiled in a close
helix. In addition to these 4 long tentacles there are 24 to 35 short, solid tentacles, not one-
fourth as long as the large ones. These short tentacles are tightly coiled, their axial cells are
chordate and they are more like cirri than tentacles. There are no ocelli or other marginal
bodies. The velum is narrow. There are 4 straight, narrow radial-canals and a slender
circular vessel. The manubrium is pynform to conical and about halt as long as the depth
of the bell-cavity. The mouth is a simple, cruciform opening. The gonads are developed in
the ectoderm on the interradial sides of the manubrium and the ova are large and con-
spicuous. The entoderm of the manubrium and tentacle-bulbs in the female (plate 6, fig. 5)
is green, but in the males light-brown (plate 6, fig. 4).

This medusa is very abundant at Tortugas, Florida, but is not so common in the Bahamas.
An apparently similar form was found by Agassiz and Mayer in the FIJI Islands, South
Pacific, although the Pacific form was duller in color than is usual in Atlantic specimens.

Genus HETEROTIARA Maas, 1905.
Hftnotiara, MAAS, 1905, Craspedoten Medusen der Siboga Exped., Monog. to, p. 19.

The type species is Heterotiara anonyma Maas, from the Malay Archipelago. Only two
imperfect specimens were found by the Siboga Expedition.

GENERIC CHARACTERS.

Tiarinae with 8 marginal tentacles (4 radial, 4 interradial). The ring-canal gives rise to
one (or more ?) blindly-ending, centripetal diverticula. Gonads ( ?)

Heterotiara anonyma Maas.
Hfterotiar/2 <mon\rna, MAAS, 1905, Craspedoten Medusen der Siboga Exped., Monog. 10, p. 19, taf. 3, figs. I9~2r.

Bell 12 to 1 6 mm. high and slightly more than half as wide. Oval, dome-like, with very
thick, gelatinous walls. 8 hollow marginal tentacles arise from the ring-canal and extend
through the gelatinous sides of the bell so as to reach the exterior at a slight distance above the
margin. The basal bulbs of these tentacles are small, and each one has a short, blunt, hollow.

108

MEDUS.E OF THK WORLD.

spur-like projection extending outward. There are no ocelli. All ot the tentacles were broken
off short and their normal length is thus unknown.

The ring-canal gives off a short, straight, blindly-ending diverticulum in one ot the 4
interradii above the base ot one of the interradial tentacles. 4 wide radial-canals as in other
Tiarinae. They are straight-edged and lack the "glandular-pouches" seen in many Tiarmae.

The circular muscles ot the subumbrella are very easily seen near the edges ot the radial-
canals and there are 4 interradial folds in the muscles which present the superficial appearance
of radial-canals, but they are merely radial muscle furrows, in no way to be confused with the
radial-canals.

The manubrium lacks a peduncle and is about halt as long as the depth ot the bell-cavity.
It is 4-sided at its base. The stomach part is urn-shaped, and thereare4toldedhps. Gonads( ?)
The tentacle-bulbs bear dense entodermal pigment granules. Color ( ?)

Two specimens were found by the Siboga expedition in the Malay Archipelago in vertical
nets drawn from 500 fathoms depth to the surface, in lat. o° 17.6' S., long. 129° 14.5' E.

It is remarkable that in each of these specimens there was but one interradial diverticulum
from the ring-canal. 4 interradial swellings are figured by Maas upon the sides of the
stomach, but he does not mention gonads. Apparently both of his specimens were immature.

FIG. 57. — " Perigonimus vestutus" Hydroid and recently

liberated medusa.
FIG. 58. — "Perigonimus m'mutus" Hydroid and young

medusa.
Above figures after Allman, in Ray Society, 1871-71.

Genus STOMOTOCA L. Agassiz, 1862.

( ?) Glatide bcroe, SLABBER, 1775, Physikal. Belustig., p. 46.

Saphenia, FORBES (non Eschscholtz), 1848, British Naked-eyed Medusa-, p. 25, plate I, fig. 4.— McCRAnv, 1 857, Gymn. Charles-
ton Harbor, p. 27.

Stomotoca, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347. — A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S.,
vol. 4, p. 347 (foot-note). — AGASSIZ, A., 1865, North Amer. Acal., p. 168.

Dinema, VAN BENEDEN, 1867, Mem. Acad. Roy. des Sci. Belgique, torn. 36, par. i, p. 127.

Amphinema+ Stomotoca, HAECKEL, 1879, Syst. der Medusen, pp. 49, 51.

Codonorchis, HAECKEL, 1879, Ibid., p. <;i.

Dinemaiella, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 151.

Stomotoca, VANHOFFEN, 1891, Zool. Anzeiger, p. 443. — MAAS, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23,
No. I, p. II.

This genus was founded by L. Agassiz, 1862; the oldest species is possibly Stomotoca
slabbcri of the northern coasts of Europe. This species was first described by Slabber, i/75>
under the name of Gladde beroc, and the hydroid and young medusa were described by Van
Beneden, 1867, as Dinema slabberi. Slabber's description is, however, so unsatisfactory that
we can not be certain that this medusa is actually a Stomotoca, and can not accept it as the
type of the genus. Stomotoca Jincma (Oceania Jincma} Peron and Lesueur may be taken as
the type of the genus. The genus Saphenia Eschscholtz belongs to the Eucopidae.

PLATE 10.

Fig. i. Dissonrtna turnJa. Tortugas, Florida, May 13, 1905. An abnormal

specimen lacking ocelli and marginal cirri.
Fig. 2. Stoniotoi'a dinema, female. Agassiz Laboratory, Newport, Rhode

Island, July, 1892.
Fig. 3. Stomotoca dinema, male. Agassiz Laboratory, Newport, Rhode

Island, September 13, 1896.
Fig. 4. Stomotoca dinema, male. Agassiz Laboratory, Newport, Rhode

Island, July, 1896.
Figs. 5 and 6. Stomotoca rugosa. Agassiz Laboratory, Newport, Rhode

Island, August 2 to 16, 1892.

Drawn from life, by the author.

PLATE 10

.\\THOMKin s K MMMOTOCA.

10!)

FIG. 59. — Perigonimus "serpens," after Alltnan, in Ray Socjety, 1871-72.

GENERIC CHARACTERS.

Tiarina; with 2 long, diamcrrically opposed tentacles, and with more or less numerous,
rudimentary tentacles. The basal bulbs of the long tentacles are hollow. The 4 interradial
gonads are complexly folded and are found in the ectoderm of the sides of the stomach. The 4
radial-canals are broad and flat and there is a well-developed circular canal. The hydroid
is Pengonimus.

There are a numberof species
of Pengonimus which probably
produce tree f.S'/omo/o, <i i medns.i-.

'.i/L \.1ia ('"'/' but the medusa is as yet unknown.

Such are: Perigontmui .\,lni,-t,/iii
Motz-Kossowska, 1905, Archiv.
Zool. Exper.. ser. 4, tome 3, p. 72,
fig. vi; a red-colored Mediterra-
nean hydroid which grows upon
Membranipora and is distin-
guished by a cup-like expansion
of perisarc at the base of each
hydranth. The clavate hydranths
have about 10 tentacles. Another
Mediterranean species is P. nnpol-
itiinns of Margin, 1904, Mirth.
Zool. Sta. Neapel, Bd. 16, p. 571,
taf. 22, fig. 25; also P. steinaehi
Jickeli, 1883, Morphol. Jahrb..
Leipzig, Bd. 8, p. 617, taf. 27, figs. 1-9, from Trieste, Adriatic. We present figures of various
species of Periganimus which produce free medusae, but in which the sexually mature
medusa is undetermined. Perigonimus mitnrcticus (Hickson and Gravely, 1907, National
Antarctic Expedition of 1901 04, vol. 3, Nat. Hist., Hydroid Zoophytes, p. 4, plates I and 4)
is found attached to the stems of Halecium in depths of o to 130 fathoms in McMurdo Bay.
South Victoria Land, Antarctic. It produces fixed gonophores, and is the only form of
Perigonimus known from Polar seas.

Stomotoca dinema L. Agassiz.

Plate 9, figs. 8 to 10; plate 10, figs. I tci 4.

Oceania dinema, PF.ROX KT Ltsi'ti R, 1809, Ann. Ju Museum d'Hist. Nat., tome 14, p. 346.— EsCHSCHOLTZ, iSi'j. s i. 1, t

Acalephen, p. 98.

Suf'henia dinema, FORBES, 1848, British Nakcd-eved MeJus.e, p. 25, plate 2, fig. 4 (Kiel. svn.).
tiaphenia titania, Gosst, 1853, Naturalists' Rambles, Devonshire Coast, p. 5X7, plate 26, figs. 7-9.
Stomotoca dinema, AC.ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.
Amphintma titania, HAF.CKEL, 1879, Syst. der Medusen, p. 50.— BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 482; Ibid.,

1905, tome 13, p. 131 (all papers to 1850).

Amphinema dinema, BROWNE, 1896, Proc. Zool. Soc. London, p. 475.
American variety = Stomotoca apicata, L. AGASSIZ.
Male:

Saphenia apifata, McCRADV, 1857, Gymn. Charleston Harbor, p. 129, plate 8, figs. 2, 3.

Stomotoca aficata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347. — AGASSIZ, A., 1865, North Ainer. Acal., p. 168.
Amphinema apicatum, HAECKEL, 1879, Syst. der Medusen, p. 50.
Female:

Dinemalel/a cavosa, FF.WKHS, 1 88 1, Bull. Mus. Conip. Zool. at Harvard College, vnl. 8, p. i?i, plate n, figs. 2. 3; plate 4, fig.

3; 1884, Amer. Naturalist, vol. 19, p. UK, I fig.

Slomotoca apicata, MAYER, 1900, Bull. Mus. Comp. Zool. Harvard College, vol. 37, p. 3, plate •. tigs. 3, 4. H
Bull. U.S. Bureau of Fisheries, vol. 24, p. 3?, 2 figs. (Figure .if male is X. apicata, but that of "female" is
NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 371, fig. 77.

r, 1904,
is S.rugoia.)—

EUROPEAN MEDUSA.

Bell ^ mm. long and 2 mm. wide, with a narrow, elongate, sharp-pointed apex. 2 long

tentacles, 2 to 10 times as long as the bell-height. About 24 small tentacle-bulbs. Time aie

no ectodermal ocelli. Stomach oval, about half as long as the depth of the bell-ca\it\ and

square in cross-section. 4 well-developed, lanceolate lips. 4 simple, slender radial-canals and

9

110

MEDUS.E OF THE WOULD.

ring-canal. The gonads are 8 adradial, transversely folded, ectodermal regions on the sides of
the stomach. Stomach yellowish-brown or green, tentacle-bulbs crimson or purplish.
Found off the coasts of Great Britain. Hydroid unknown.

I have seen medusae oft" the coast of Cornwall, Eng-
land, which were identical with Stoniotoca apicata of
America.

AMERICAN MEDUSA.

In the adult American medusa the bell is about 4
mm. high and 2 mm. wide. There is a large apical
projection which is hollow in the female, but usually solid
in the male medusae. The bell-walls are thin and the
sides vertical below, but slopinginward above to a pointed
apex. There are 2 long, diametrically opposed ten-
tacles, with large, tapering, hollow basal bulbs. There
are no ocelli. These long tentacles are highly contractile,
but are usually about as long as the bell-height. Their
shafts are covered with small wart-like clusters of nema-
tocysts. In addition to the 2 long tentacles, there are 6
or more small rudimentary tentacle-bulbs upon the bell-

FIG. 60. — Stoniotoca dinema, male. From life, by
the author. Mousehole, Cornwall,
England, October 28, 1907.

Synopsis of the Species of Stoniotoca.

S. dinema
Haeckel.

S. "apicata"
L. Agassiz.

S. atra
L. Agassiz.

S. octaedra.

S. rugosa

Mayer.

S.pterophylla
Haeckel.

S. divisa
Maas.

Size of bell
in mm.

3 long, 2 wide.
Apex solid.

4 high, 2 wide
with solid
apex in male
and hollow
apex in
female.

25 high, 22

wide. Apex
solid.

2.5 high, 2
wide. Apex
solid.

5 high, 3 wide.
Apex large,
solid. Sides
thin.

30 wide, 12 high.
Small apical
projection.
Flaring sides.
Thick gelati-
nous substance

30 wide, 20
high. As in
S. ptero-
phylla.

Length of
tentacles

6 to 30.

4

io±

5±

30±

20 ±

AsinS.

pterophylla.

in terms

of bell-ra-

dius (r).

Number of
rudimen-

24

6

80

6

'4

60 to 80.

As in S.
pterophylla.

tary ten-
tacle-

bulbs.

Ectodermal,
abaxial,

None.

None.

None.

8, orange-
colored.

None.

None.

None.

tentacular

ocelli.

Shape of
manu-
brium.

Oval, half as
long as
depth of
bell-cavity.

Flask-shaped.
4 folded,
cruciform
lips.

Half as long as
depth of
bell-cavity.
Gonads
folded as in

As in S. api-
cata.

As in S. atra.

As in S.atra.

As in S.
pterophylla.

Turris.

Color.

Stomach

In male, stom-

Mouth dull-

Stomach and

Stomach, gon-

Stomach and

Gonads

yellowish-
brown to
green. Ten-
tacle-bulbs
purple or
crimson.

ach green
and tenta-
cle-bulbs
purple. In
female, dull
milky-yellow.

yellow,
gonads very
dark-brown,
nearly black,
tentacles
light-brown.

tentacles
green, with
orange cen-
ter in stom-
ach.

ads, and ten-
tacle-bulbs
brick-red
to nearly
black.

tentacle-bulbs
brown. Radial-
canals and
gonads milky-
white.

orange to
cinnabar-
red. Stom-
ach and ten-
tacle-bulbs
yellow.

Where

found.

Coast of Great
Britain.

Atlantic coast
of United
States.
Probably

Port Town-
send, Wash-
ington.
Pacific coast

Tortugas and
the Baha-
hamas, in
summer.

Woods Hole to
Tortugas,
Atlantic
coast of

Bahamas, Tortu-
gas, West
Indies.

Pacific coast
of Central
America.

identical

of America.

United

with S. di-

States.

nema.

PLATE 1 1 .

Figs, i and 2. Stotnotoca rugosa, young and mature. Newport, Rhode
Island, August 4, 1892.

Fig. 3. Young medusa of Perigonimus jonesii. Agassiz Laboratory, New-
port, Rhode Island, August 22, 1896.

Fig. 4. Young medusa of Perigonimus jonesii. Agassiz Laboratory, New-
port. Rhode Island, July, 1896.

Fig. 5. Stotnotoca octaedra, young medusa. Tortugas, Florida, July 17,
1898.

Fig. 6. Stomotoca octaedra, Tortugas, Florida, June, 1897.

Fig. 7. Pandca violacca. Aboral view of gonads of the medusa shown in
figure i, plate 12.

Drawn from life, by the author.

r

\ vrin IM Knrs.K- -ST< >M< >rt ><-.\. Ill

margin. These lack ocelli. The 4 radial-canals are wide and flat. The boundaries of the
radial-canals and of the ling-canal are often bluntly serrated. The manuhrium is flask-
shaped; there is no peduncle and the 4 lips are cruciform, crenulated, and curve slightly
upward. The ectoderm of the upper part of the manuhrium on both sides of the 4 ladial-
canals is thrown into interradial folds or convolutions and the gonads are developed in
this region.

In the male the color of the manuhrium varies from intense green to translucent ocher-
yellow or cream-color. The basal bulbs of the tentacles in the male vary from faint to deep
purple. In the female the manubrium is usually translucent ocher-\ t -\\n\\ or cream-colored,
but occasionally straw-colored or faintly green. The tentacle-bulbs of the female are usualU
translucent milky or ocher, but occasionally one is found having faintly purple tentacle-bulbs.
These sexually dimorphic color peculiarities are seen in the smallest and youngest medusae
as well as in adults. In very young medusae the apical projection is small or absent. There are
2 tentacles and only 2 rudimentary tentacle-bulbs.

This medusa is common at Newport, Rhode Island, from |ulv 15 to September. It is
not common at Tortugas, Florida, and is not seen north of Cape Cod, Massachusetts. On
October 28, 1907, I found two male medusae of this species oft" Mousehole, Mounts Bay,
Cornwall, England.

This species furnishes that which is probably the most remarkable instance of sexual
dichromatism to be met with among the hydromedusae, the color differences affecting parts
other than the gonads.

The two males found off the coast of Cornwall, Kngland, were each 2.2 mm. high and
similar in all respects to the typical American form; with intense green manuhrium and
purple tentacle-bulbs, without ocelli. The bell-apex was solid and gelatinous. 1 believe that
S. apicata of America. IS identical with S. Jlni'inn of Europe. The European medusa may
have more rudimentary tentacle-bulbs, hut these are very variable in number in the Ameri-
can form.

Stomotoca atra Agassiz.

Stottiotora atrn, AGASSI/, I... i862f Cont. Nat. Hist. U. S.f vol. 4, p. 347.- A<.-\SM/, A., iM>^. N^rth AIIMT. \<,il., p. it>^, figs.
271-27;. H^H-KH., 1X71), Sysl. ilcr Meilusen, p. 53.

Bell 20 to 25 mm. high, 20 to 22 mm. wide. Bell-shaped, higher than a hemisphere.
2 long, highly contractile tentacles, and about 80 small rudimentary tentacle-bulbs. Stomach
about half as long as the depth of the bell-cavity. Swollen, 4-sided, and mounted upon a
peduncle. 4 lips. The gonads a re 8 ad radial, linear cross-foldings upon the sides of the Stomach,
each gonad consisting of I 2 to 15 folds as in Turns. The mouth is dull yellow. Gonads dark-
brown to almost black. Tentacles light-brown.

Found by A. Agassiz in great numbers in Port Townsend, Washington, Pacific coast of
the United States; from |une until September.

Stomotoca octaedra.
Plate 1 1, ti);s. <; ami f>.

Cotlunorchis octnetlrus, HAFCKK.L, 1879, Syst. Jrr MrtliiM-n. p. 51.

Sloniolocn austrtilis, MAYKR, 1900, Bull. Mus. (.'nrn|'. /.mil. .it Harvard College, vol. 37, p. 32, plate I, lii;. i; lintl., 1904,
Mrin. Nat. Sci. Brooklvn Inst. Museum, vol. i, p. 9, platr i, fig. 9.

The following description is derived from studies of specimens found in the Baham.is.
and at Tortugas, Florida.

Immature nirJ/isn (plate 11, fig. 6). — Bell about 2.5 mm. high and with well-developed,
sharp-pointed, apical projection upon aboral side. Bell-walls thin.' There are 2 radially
situated, diametrically opposed tentacles, each about as long as the bell-height and with
long hollow, tapering basal bulbs. There is a single, ectodermal, orange-colored ocellus upon
the outer side of each tentacle-bulb near its point of origin from the umbrella. In addition to
the 2 long tentacles there are 2 short, rudimentary tentacle-bulbs go0 from the long tentacles.
Each of these rudimentary tentacle-bulbs has an ectodermal, orange-colored ocellus upon its
outer side, and there are 4 other ocelli upon the bell-margin in interradial positions. There
are thus 8 ocelli in all, 4 on the tentacle-bulbs and 4 interradial ones on the bell-margin. 'I he

112 MKDUS.E OF THK WOULD.

velum is well developed. There are 4 straight-edged, wide radial-canals and a broad circular
vessel. The manubrium is urn-shaped, quite wide, and with 4 cruciform, recurved lips. The
mouth is at a point about half-way between the inner apex of the bell-cavity and the velar
opening. The gonads are found in complexly folded regions upon the 4 interradial sides of
the stomach. The manubrium and tentacle-bulbs are yellow or greenish-yellow. The ento-
dermal core of the stomach is often orange.

When about I mm. high (plate 1 1, fig. 5) the bell has a small apical projection. There
are 2 large tentacles and 2 rudimentary tentacle-bulbs, each with an orange ocellus. There
are no other ocelli upon the bell-margin. The manubrium is slender and urn-shaped, without
gonads and with 4 simple, cruciform lips.

The young of this medusa are common at Tortugas, Florida, and in the Bahamas through-
out the summer. Although the medusa has usually but 2 long tentacles, occasionally one is
seen with 4 long, equally developed, radially situated tentacles. The mature medusa has not
been seen and we must remain in doubt concerning its generic position, for it may be a young
Turn's. The presence of ocelli upon the tentacle-bulbs distinguishes it from all other Ameri-
can species of Stornotoca and is a character commonly seen in Turns.

I believe this medusa to be identical with Haeckel's " Codonorchis octaedrus," which he
obtained offthe Atlantic coast of France. Haeckel states that this medusa had 2 well-developed
tentacles and 10 tentacle-bulbs (2 perradial and 8 adradial). He described the ocelli as
brownish-red. Apparently he found but a single specimen and the interradial tentacle-bulbs
may have failed to develop. His medusa is described as having folded gonads, and is said to
be 4 mm. high and 2.5 wide.

Stomotoca rugosa Mayer.
Plate 10, figs. 5 and 6; plate 11, figs, i and 2.

Slomoloca afirala, FEWKF.S, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 152, plate 2, figs, i, 4, 9.— HARC.ITT,
1901, American Naturalist, vol. 35, p. 581, fig. 40; 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 35, i fig. (errone-
ously labeled Stomotoca apicata, female). — RITTENHOUSE, 1907, Proc. Boston Soc. Nat. Hist., vol. 33, pp. 440, 445,
452, 456 (embryology).

Stomotoca rugosa, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 4, plate 2, fig. 5; Ihitl., p. 32.

Amphinema u[titatuin, BROOKS, 1883, Studies Johns Hopkins Biol. Lab., vol. 2, p. 473.

Bell 5 mm. high and 3 mm. broad. Solid, apical projection, in some individuals short and
blunt, in others long and slender. Bell-walls of moderate thickness, and bell itself, exclusive
of the apical projection, somewhat higher than a hemisphere, with relatively vertical, straight
sides. There are 2 long, diametrically opposed tentacles, which are of equal length, and when
stretched are fully ten times as long as the bell-height. They are, however, highly contractile
and may become not more than one-tenth as long as when fully expanded. The basal bulbs
of these long tentacles are large, hollow, and tapering and there are no ocelli.

In addition to the 2 long tentacles there are 14 small, permanently rudimentary tentacles,
2 at the bases of 2 of the radial-canals and 3 in each interradial quadrant. These lack ocelli.
The velum is well developed. There are 4 wide radial-canals and a ring-canal, all with
jagged edges. The manubrium is quadratic and flask-shaped and the mouth in old medusae
is about at the level of the velar opening. The mouth is cruciform and there are 4 prominent,
recurved, crenulated lips. The mature sexual products are found in the ectoderm of the
adradial walls of the stomach on both sides of the places of entrance of each of the 4 radial-
canals where the surface is thrown into 8 series of complex folds and ridges, a double ridge of
folds in each interradius.

The gelatinous substance of the bell is transparent, but the entoderm of the tentacle-bulbs
and stomach is brick-red, often streaked with sooty brown. The radial and circular canals are
faint red in color. Specimens from Tortugas, Florida, often show black streaks through the
brick-red color of the stomach and tentacle-bulbs, and medusae which have been confined in
aquaria for some days often become wholly black in these parts.

Hydrotd and \onng medusa. — Professor W. K. Brooks, 1885, describes the hydroid.
It is a Pi-rigfjniiniis, very much like P. niiniitiis Allman, 1871 (Monog. Tubularian Hydroids,
p. 324, plate xi, figs. 4 -6). It was found at Beaufort, North Carolina, growing upon the lower

ANTHOMKIH'S.K— STOMOTOfA. 113

surface of the shells of Limulus, fastened to the sand-tubes of Sabfllarin. The stems are simple
and unbranched and are only about 0.2 mm. in height. They are covered for about two-
thirds of their length by a delicate, closely-adherent film of perisarc to which foreign particles
become attached. The stomach occupies about one-fourth or one-fifth of the length of the
stem from which it is separated by a slight constriction. Each polypite has 10 tentacles which
point alternately backwards and forwards, those pointing forwards being a little longer than
the others. The medusae are attached by very short peduncles to the sides of the stems. When
the medusa is set free it is about 0.5 mm. in height and there is no trace of the apical projection,
which develops in about eight days.

In an abnormal individual medusa of this species found at Newport, Rhode Island, in
July, 1892, there were 4 long tentacles, i at the base of each of the 4 radial-canals. This
medusa was maintained alive in an aquarium for more than a month. When first found it had
only 2 long tentacles which were diametrically opposite one another at the bases of i of the
radial-canals. The other 2 tentacles developed later, after the first pair had attained their full
length. The medusa had then 4 radially placed tentacles and 12 rudimentary tentacle-buds.
This variation is interesting, as it illustrates the close relationship between Stiiniutrnu and
Turns.

Stomotoca ruorjsa is common on the southern coast of New England in summer. It is
found all along the coast to southern Florida, but is not very common at Tortugas or among tin-
Bahama Islands. It has never been seen north of Cape Cod, Massachusetts.

Rittenhouse, 1907, finds that the eggs of this medusa are laid between 5'' to 5'' 30"' in the
morning. The egg is chalky-white. The entoderm is formed by cellular ingression. The
planula settles down upon its side and becomes a branched hydrorhi/a from which the p<>!\ •-
piles bud out.

Stomotoca pterophylla Haeckel.

Plate 2<), figs. 5 to ^; pl.Ue 30, fig. 7.

Xtomotoca ptfroplnlla, HAKCKF.I., 1879, Syst. der Medusen, p. 52, taf. 4, fig. 10.
Xiumolorti persphyHa, F I:\VKKS, |SX<), Report Comrmss. Fish and Fisheries for 1X86, p. 524.

Adult medusa. — Bell conical, with widely flaring sides and small, sharply-pointed apical
projection. It is about 20 to jo mm. in diameter and 10 to 12 mm. in height. The gelatinous
substance is very thick at the aboral pole, but becomes thin at the bell-margin. There are 2
long, tapering, marginal tentacles which are situated at the bases of 2 of the radial-canals, 1 80°
apart. When expanded these tentacles are full}- 10 times longer than the bell-diameter. In
addition to the long tentacles, there are about 60 to 80 small rudimentary tentacle-bulbs, 15 to
20 in each quadrant. The 4 radial-canals are wide, ribbon-like, and flat, with smooth, simple,
outer edges. The ring-canal is narrow and its upper edge is smooth. There is a very wide,
conical peduncle which extends about to the level ot the velar opening. The gastric part of the
manubi mm is large and swollen and lies mainly outside ot the bell-cavity. There are 4 promi-
nent, complexly crenulated, lanceolate lips. The gonads occupy 8 adradially situated, longitu-
dinal swellings upon the sides ot the stomach (plate 29, fig. 4). Each gonad consists ot a io\\
of swollen, leaf-like ridges which trend in a transverse direction. The stomach and tentacle-
bulbs are brown. Radial-canals milky. There are no ectodermal ocelli.

This species is found in the West Indies and the warmer parts of the Gulf Stream. I
found many specimens among the Bahamas and at the Tortugas during the spring and early
summer ot 1907.

It is closely related to Stomotoca Jivisa, described by Maas, from the west coast of Mexico
(Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. n, tat. i. tigs, i i<j, 180,7). 'Hlt
its rich-brown entoderm distinguishes it from the Pacific species.

Haeckel describes this medusa from preserved specimens, and this may account for
certain errors in his description and figure. A small apical projection appears to be con-
stantly present and there are no "ocelli." The radial and circular muscles of the velum are
very powerfully developed. In extreme states ot contraction of the bell the peduncle mav
extend beyond the velar opening or tall short of reaching it. The long tentacles are higlih
contractile and when expanded are reduced to mere thread-like filaments.

Ill MKDrS.E OF THE WORLD.

Stomotoca divisa Maas.

Siomoioca Jk'isa, MAAS, 1897, Mem. Museum Comp. Zool. at Harvard College, vol. 23, p. u, taf. i, fign. 1-9.

Bell 20 to 30 mm. wide, 15 to 20 mm. high. Upper part of bell thick, solid, and dome-
like, and separated by an annular furrow from the thin-walled, marginal part of bell. This

furrow may be due to contraction ( ?) A
very small, pointed apical projection arises
sharply from the aboral surface of the
evenly rounded dome of the bell. There
are 2 tapering tentacles about as long as the
bell-height. These appear to have some-
what more swollen basal bulbs than are seen
in S. pterophylla. There are no ocelli. There
are a large number of rudimentary tentacle-
bulbs as in S. pterophylla. The bell is trans-
parent with yellow entoderm, and with
orange to cinnabar-red gonads.

IMI.. 61. — Stomotoca Jivisa, after Maas, in Mem. Museum Comp. It IS found in the Bay oi Panama,

Zool. at Harvard College, showing different statesof Pacjfic coast of Central America, ill March,
contraction of the bell. -.......,- ... .

It is distinguished from the closely allied

.S\ pterophylla of the West Indies by its brilliant coloration, S. pterophylla being constantly
dul I -brown.

Perigonimus jonesii Osborn and Hargitt.
Plate n, figs. 3 and 4.

Perigonimus jonesii, OSBORN and HARGITT, 1894, Amer. Naturalist, vol. 28, p. 27, figs. 1-12. — HARGITT, 1895, Mittheil. Zool.
Station Neapel, Bd. u, p. 479; 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 33, I fig.; 1901, American Naturalist,
vol. 35, p. 308, fig. 4; p. 579, fig. 37. — NUTTING, 1901, Bull. U. S. Fish Commission, vol. 19, pp. 331, 372, fig. 80.

} oung medusa. — None of the specimens yet seen were mature. Bell of largest 2 mm.
in height and side walls extremely thin and flexible. There is a very small, dome-shaped
apical projection. There are 2 well-developed, diametrically opposed tentacles, which are
situated at the bases of 2 of the radial-canals. These tentacles are of unequal length and
are at times carried curled in a close helix and at other times are extended to their full length,
in which case one of them becomes about as long as the bell-height, while the other attains to
about twice this length. The basal bulbs of these tentacles are long, conical, and hollow
and have no ocelli, and the shafts of the tentacles are thickly covered with nematocyst-cells.
In addition to these well-developed tentacles, there are 2 small tentacle-bulbs, situated at
the bases of the 2 radial-canals 90° away from the long tentacles. There are no ocelli. The
velum is wide and flexible. There are 4 straight, slender radial-canals and a narrow, circular

' O

tube. The manubnum is short and simple, with a wide base and 4 simple, cruciform lips.

The ectoderm of the manubrium and tentacle-bulbs is of a dull ocher-yellow.

This medusa is very rare; only 3 specimens have been seen by me during 3 summers' study.
All of my specimens were found in Newport Harbor, Rhode Island, during July and August.

Hydroid. — The hydroid has been described by Osborn and Hargitt, 1894, from Cold
Spring Harbor, Long Island, New York, where it is found very commonly upon the abdomen
and upon the walking legs of the spider-crab (Libinia emarginata). It is a Pengoniiniis.
The stems arise from a creeping hydrorhiza and branch luxuriantly. The oldest polypite is
found at the distal end of the stem. The stems are covered with a thick gelatinous pensarc
which extends up the stems to the level of the tentacles, to which it is fastened. There are
normally 16 tentacles in a single circlet around the base of the hypostome. These assume an
alternately reflected position. The medusa-buds arise in clusters from near the center of the
stems. They are covered with the thick perisarc and each one is attached to the stem by
means of a well-developed peduncle. When set free the medusa has 2 diametrically opposed
tentacles which are usually carried turned inward into the bell-cavity. The hydroid is flesh-
colored.

The medusa can at once be distinguished from S. apictita by the extreme tenuity and
flexibility of the bell-walls. The bell is also higher and narrower than in S. apicata The

ANTIKiMKDIS.K ST< iM( iT( i( 'A, DISSi i\KM \ . II.")

medusa swims by the aid ot a rhythmical series ot wave-like contractions which travel one
after another up the very flexible sides ot the hell, from margin to apex. The efficiency ot
these movements is greatly enhanced In the cooperation of the powerful velum.

The 4ully-developed medusa is unknown and it-is possible that this medusa may not be
Stomotoca. It can not be referred to any Turns known upon our coast, and the tact that the
2 long tentacles are of unequal length, one being considerably longer than the other, separates
it at once from any other known species of Sttjtnotrjici. At present we place it provisionally
in the genus Strjinotoca to which its hydroid appears to refer it. Were it not for the absence
ot ocelli upon the tentacle-bulbs I would be inclined to suspect that this medusa might prme
to he the young ot Turns pileato.

(Stomotoca?) Perigonimus cidaritis Weismann.

Pfrigottiniiis cidaritisi WUSMANN, 1885, Entstehung Sexualzellen bei Hyiimmciluscn, pp. 117, 218, taf. 12. figs. 10, 11.

The stems of the hydroid are about 90 mm. high and arise thickly one by the side of the
other from a root-like hydrorhiza. The stems are often simple and unbranched, but tlu\
commonly branch dichotombusly one or two times, the branches being set off at acute angles
one with another. The hydranths have a circlet ot about 20 tentacles. Perisarc thin and
yellowish. The medusa-buds are borne upon pedicels in a zone at the bases ot the hydranths.
When set free the medusa is spherical, with 4 marginal tentacles and 4 radial-canals. The
manubrium has 4 short, knobbed oral tentacles. The whole hydroid stock is invested with
a thin layer of slimy mud, leaving only the mouth-ends of the hydroids projecting, and some-
times even those are covered, leaving only the tentacle tips tree.

Found at Naples, Italy. Medusa set tree in March.

The sex-cells originate in the ectoderm ot the proximal part of the manubrium of the
medusa-bud and develop in their place of origin, becoming mature in the tree medusa. The
mature medusa has not been determined.

Genus DISSONEMA Haeckel, 1870.

l)i«onrma, HAF.I KM, 1X7.), Svst. dcr Mrdusen, p. 126.— MAYF.R, 1900, Bull. Mus. C'oinp. /.mil. .11 H.ir\.ir,l (.'iill'-i'r , \nl. ;-,
p. 44. — BK;F.UHV, H. B., 1909, Bull. MiiM'um of Comp. Znol. ;it Harvanl Colli-iv.

This genus was founded by Haeckel, \$~(), for Dissotit-itiii .\ii[>lii'n,-llii, of the- coast <>l
Australia.

GENERIC CHARACTERS.

Tiarinas with two hollow, diametrically opposite tentacles. The gonads extend from
the manubrium outward along the canals. Without marginal sense-clubs. Cirri aie some-
times present. With abaxial, ectodennal ocelli upon the bases ot the tentacles.

Rigelow, 1408, finds that the adradial gonads begin to develop upon the sides of the
manubrium and that afterwards they extend outward over the radial-canals. The hollow
tentacles, abaxial ectodermal ocelli, rudimentary tentacle-bulbs, and hollow bell-apeK are
all characteristic of the Tiarinae., and it appears that this genus must be removed trom the
Thaumantiad;e, in which Haeckel placed it.

J)is.friiii-niii bears the same relation to St»in<tt't,<i that .\<//;<//MM does to Bougainvillta.
The hydroid is unknown.

Dissonema saphenella Haeckel.

/>;\w,'M w<; niplifntlla, HAFI KKI, 1879, Svst. tier Mcduscn, p. 126, t:tf. 8, rl^. }.

Bell pyriform, with thick, solid apex and thin, bulging sides. 6mm. high, 4 mm. uulc.
2 well-developed tentacles, several times as long as the bell-height. These tentacles have
large, swollen, conical basal bulbs. There are 2 rudimentary tenta'cle-bulbs yo° apart trom
the long tentacles. There is an abaxial (ectodermal ?) "ocellus" ot large si/e upon each ot
the 4 tentacle-bulbs. There are no marginal clubs, cirri, or other appendages.

The velum is well-developed and there are 4 straight, narrow radial-canals and a ring-
canal. The manubrium is cylindrical, half as long as the depth of the bell-cavity and pro-
vided with 4 short, crinkled lips. The 4 folded, spindle-shaped »onads are developed upon
the middle three-fourths of the radial-canals. Color ( ?) Coast ot Australia. (See tig. '

Ill)

MEDUS.E OF THE WORLD.

Dissonema turrida Mayer.

Plate 10, fig. l; plate 22, fig. I.

Dissonema turriJa, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 44, plate 2, figs. 3, 4; 1904, Mem.
Nat. Sci. Brooklyn Institute Museum, vol. i, p. 8, plate 2, fig. 10.

Adult medusa. — Bell bluntly cone-shaped with a hollow apical projection. It is 4 to 7
mm. high and the side walls are thin and flexible. There are 2 long, hollow, gradually tapering

tentacles which are quite contractile, but
when expanded are 3 to 4 times as long as
the bell-height. Each of these tentacles has
a minute, red ocellus in the ectoderm of its
outer side near the bell-margin. In addition
to the two large tentacles there are 12 to
14 small, solid cirri which arise separately,
at equal intervals, from the bell-margin.
Each cirrus has a red ocellus in the ecto-
derm of its outer (abaxial) side at the bell-
margin. The velum is well developed.
The circular vessel is narrow, but the 4
radial-canals are broad, and the 4 convo-
luted, bag-like gonads occupy three-quar-
ters of their length and also lie upon the
adradial sides of the manubrium. In the
female each gonad contains 4 to 7 large eggs
which project prominently over the surface
of the organ. The manubrium is pyritorm
and the mouth projects beyond the velar
opening. The stomach-walls are thin and
flexible, and the mouth is surrounded by
large recurved lips with crinkled edges. 1 he
entoderm of the manubrium and tentacles
is a delicate green and the genital organs
and circular canal are tinged with green or
pink. The ocelli are red.

In young medusae there are but 2 large
tentacles and 2 cirri. There are 8 ocelli.
The 2 long tentacles have tapering, hollow
basal bulbs, but their main shafts are at
first solid, though later they become hollow.
This medusa is common on the surface
in the Bahamas and at Tortugas, Florida,
in summer.

Dr. H. B. Bigelow shows as a result of
sectioning the medusa of Dissonema that the
gonads begin to develop upon the adradial
sides of the manubrium and later extend
outward down the radial-canals. This fact,
taken in connection with the abaxial ectodermal ocelli, the hollow tentacles, bell-apex, rudi-
mentary tentacles and general form of the medusa, make it evident that Dissonema is one
of the Tiarinne.

Genus PANDEA Lesson, 1843.

Pan/tea, LESSON, 1837, Prodrom. Monog. Meduses, No. 22 (not published); 1843, Hist. Zooph. Acal., p. 288. — ACASSIZ, L.,

1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.
Pantlepa, HAECKEL, 1879, Syst. der Medusen, p. 53.
Pandra, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 160. — HARTI.AI-B, 1892, Nachricht.

kgl. Gesell. Wissenschaft. Univ. Gottingen, p. 21. — MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 15.
Diantfa, Quov ET GAIMARD, 1827, Annal. des Sci. Naturelles, tome 10, plate 6.
.\'on Dianefa, LAMARCK, 1816, Hist. Anim. sans Vert., tome 2, p. 505.
Pandja, ZOJA, 1892, Boll. Sc. Paira, Anno 1891, Nos. 3, 4, 3 pp.; 1892, Annals and Mag. Nat. Hist., ser. 6, vol. 9, p. 409.

(Deniiroclam dohrnii gives rise to a medusa resembling Pandea ?)

Fin. 62. — Dissonema saphenella, after Haeckel, 1879.

.\\THilMKDCS.E— r.\\T)K\.

11;

The type species of PanJea is P. cnnlca Lesson, 1843. This medusa was previously
described by Ouoy and Gaimard (1827, Annal. Sci. Nat., torn. 10, p. 182, plate 6) under the
name Diantea conica. It is found in the Mediterranean. Haeckel introduced the spelling
" PanJ(ta," although Lesson's original spelling is PanJea. The genus "Diantea" was founded
by Lamarck (1816, Hist. Anim. sans Vert., torn. 2, p. 505), the type species being Diancea
tneJra, a medusa which had been previously described by Pt-ron under the generic name of
Lymnorea. Lamarck's genus Duimta also included various species of Geryonta, Oceania,
Pelagia, and Mfdusa of previous authors, and is so hopelessly involved that in my opinion
it must be dropped.

GENERIC CHARACTERS.

Tiarinae with 4 or more well-developed, marginal tentacles. There are 4 interradial,
genital ridges in the ectodermal wall of the stomach, but these gonads are only imperfectly
separated in the 4 principal radii; so that the stomach is completely encircled by the genital
organs, which are, however, better developed in the interradii than in the principal radii.

DenJroclava is possibly the hydroid of some species of PnnJea, Coins, or Turn's-, but the
mature medusa is unknown.

When set free the medusa of DenJroclava dohrnii (Weismann, 1883, pp. 26, 216, taf. 12,
fign. 6-9) has 8 marginal tentacles, 4 radial-canals, and a manubrium with 4 band-like gonads

i<;. 6j. — Pandeei conica.

A. Side view of half-grown medusa. Bell 13 mm. high. Gonads interradial.

H. Manubrium of mature medusa, with bell 21 mm. high. Gonads continent, surrounding stomach.
C. Manubrium of young medusa, showing interradial gonads. Bell (> nun. lir\;h. l-'rom lit- . I'.
the author. Zoological Station, Naples. Winter of 11)07-08.

on its interradial sides. The sexual cells originate in the ectoderm of the manubrium of the
medusa-bud and develop in the ectoderm of the free-swimming medusa, without wandering
from their place of origin. The hydroid of DenJroclava dohrnii was found by Weismann at
Naples in the summer of 1881, attached to a stock of Isis which was dredged from a depth ot
35 to 40 fathoms. The stems are about 35 mm. high and arise from a short, creeping hydro-
rhiza, which together with the stems is covered by a smooth, horny perisarc. The main stems

118

MKDUS.E OF THE WOULD.

give rise to alternate side branches. The hydranths are cluh-shaped and bear 12 to 20 short,
filiform tentacles, scattered over their whole sides. The medusa-buds arise singly from the
side branches close under the neck of the lateral hydranths. Pictet, 1893, found a very similar
or identical hydroid at Amboina, Malay Archipelago. The principal references to this clava-
like hydroid are as follows:

DenJroflava Johrnii, WEISMANN, 1893, Entstehung Sexualzellen bei Hydromedusen, Jena, pp.

26, 215, taf. 12, fign. 6-9. — DU PLESSIS, 1888, Reeueil Zool. Suisse, tome 4, p. 531 (from

Villefranclie).
DenJroclava dohrni, PICTET, 1893, Revue Suisse Zool., tome I, p. 6, plate I, figs. I, 2 (from

Amboina).

It does not seem probable that Dcndroclava dohrni produces a Pandea-like medusa, for it
is well established by Brooks that Tumtopsis nutricula comes from a DenJroclava hydroid.

Pandea conica Lesson.

Dian<ta conica, p_i OY IT GAIMARD, 1827, Annal. des Sci. Nat., tome 10, p. 182, plate 6 A, figs. 3, 4.

Pandea conica, LESSON, 1843, Hist. Zooph. Acal., p. 288. — HAECKEL, 1879, Syst. der Medusen, p. 54. — MAAS, 1904, Result.
Camp. Sci. Prince de Monaco, fasc. 28, p. 16, plate i, figs. 6, 7 (figures of gonads). — BEDOT, 1905, Revue Suisse de
Zool., tome 13, p. 141 (literature, 1827-50). — LOBIANCO, 1903, Mitth. Zool. Sta. Neapel, Bd. 16, p. 217.

Oceania setiecimcostata, KOLLIKER, 1853, Zeit. fiir wissen. Zool., Bd. 4, p. 324.

Oceania conica, GEGENBATR, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 221, taf. 7, fign. 1-3.

Bell with barrel-shaped sides, and bluntly pointed or concave apex; 21 mm. high, 10 mm.
wide. There are 8 to 24 well-developed, longitudinal rib-like ridges in the tentacular radii on
the exumbrella, and an equal number of marginal tentacles with abaxial ectodermal ocelli.
Stomach wide and short with 4 folded lips having sinuous margins. The gonads consist ot 4
crescentic, reticulated, swollen regions in the ectoderm of the 4 interradial sides of stomach,
the convexity ot the horseshoe pointing aborally. These gonads are not completely separated
in the adult, in the 4 principal radii, so that they surround the stomach on all sides.
The best figures illustrating their structure are given by Maas, 1904.

Stomach brownish. Gonads and
tentacle-bulbs brownish-red. Tentacles
yellowish-milky. Ocelli dark-purple.
Common in the Mediterranean.
This medusa was quite common
during the winter of 1907-1908 at
Naples, Italy. When young the 4 inter-
radial gonads are completely separated,
but later they fuse more or less over the
perradii, forming a complete, swollen
network which girdles the stomach.
The prominent longitudinal ridges over the exumbrella arise after the tentacles begin to
develop and extend upward from the bases of the tentacles toward the bell-apex. The table
gives a growth-record obtained by the author from specimens of this medusa observed at
Naples. (See fig. 63, page 117.)

Pandea saltatoria Lesson.

Oceania saltatoria, SARS, 1835, Beskriv og Jagtt., p. 25, plate 4, fig. loa-r.

Pandea sallatoria, LESSON, 1843, Hist. Zooph. Acal., p. 290. — HAECKEL, 1879, Syst. der Medusen, p. 54. — BEDOT, 1901, Revue
Suisse de Zool., tome 9, p. 485; Ibid., 1905, tome 13, p. 141 (all literature to 1850).

Bell 8 mm. high, 6 mm. wide, with a pointed apex and rounded sides. Exumbrella with
only 12 to 16 longitudinal lines of nettling-cells instead of about twice as many as in P. conica.
24 to 32 tentacles; twice as many as the lines of nematocysts, instead of being equal to these in
number as in P. conica. Ocelli on outer sides of tentacle-bulbs. Stomach slender, lips small
and simple. Immature (?) Color (?) Coast of Norway, Bergen.

Pandea minima von Lendenfeld.

Pand<ra minima, vox LEXDFNKELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 916, plate 42, figs. 10-12.

Bell semi-ovate, 3 mm. high, 2 mm. wide. 8 tentacles about as long as bell is high. 8
adradial lines of nettle-cells upon the exumbrella. A bundle of nerve fibers is said to extend
under each linear series of nettle-cells connecting the clusters one with another! The man-

Height of
bell in mm.

No. of

tentacles.

Condition of the gonads.

6

16 4 small interradial.

X

16 4 larger interradial.

'3

21 4 interradial nearly touching in the

perradii.

21

24

A unitary network of ridges sur-

rounding the stomach.

PLATE 12.

Fig. i. Pandea violacca, mature female. Tortugas, Florida, June 5, 1906.

Fig. 2. Turris vesicaria, young medusa. Eastport, Maine, September 19,
1898.

Fig. 3. Turris vesicaria, half-grown medusa. Agassiz Laboratory, New-
port, Rhode Island, June 24, 1893.

Fig. 4. Turris pileata, half-grown medusa. Agassiz Laboratory, Newport,
Rhode Island, July 13, 1895.

Fig. 5. Podocoryne fulgurans. Basal part of one of the marginal tentacles.

Figs. 6 to 9. Podocoryne fulgurans. Views of manubrium showing medusa-
buds. Agassiz Laboratory, Newport, Rhode Island, September,
1892.

Drawn from life, by the author.

fin

x

ANTHOMKDU.S.E — PANDEA.

II '.I

ubrium is slender and about half as long as the depth of the bell-cavity. 4 small lips. 4
longitudinal gonads on the stomach. These are narrow folds with smooth outer surfaces.
Bell light-pink, with the 8 adradial lines of nematocysts more intense in color. Stomach and
tentacle-bulbs light-brown.

Found in Sydney Harbor, New South Wales, Australia, in August and September.

Pandea violacea Agassiz and Mayer.

Plate ll, fig. 7; plate 12, fig. i.

Pandra violacea, AGASSIZ, A., and MAV>R, iX<)<l, Bull. Mus. Comp. £<»>!. at Harvard C'"llrt;i\ vol. ;2, p. i(>o. M\vtR, 1900,

Bull. Mus. Comp. Zocil. at Harvard College, vol. 37, p. 34, plate i, fig. i.
Pandea, s/>., LoBiANCo, 1903, Mitth. Zoolog. Sta. Neapel, Bil. 16, p. 217, lav. 7, fig. i.

Bell pear-shaped and 7.5 mm. high, with moderately thick
walls. There are about 32 tentacles. 8 to 12 of these are
about 3 times as long as the bell-height and 24 are small and
rudimentary. The long tentacles are all of equal length. Their
basal bulbs are long, tapering, and hollow. There are i to 3 rudi-
mentary tentacle-bulbs between each successive pair of long ten-
tacles. There are about 32 ectodermal, purple ocelli, one on tin-
outer side of each tentacle-bulb. The velum is well developed.
The manubrium is flask-shaped and quadratic in cross-section.
The outer surfaces of the adradial gonads are smooth and with-
out corrugations (plate n, fig. 7). The mouth is at the extrem-
ity of a well-developed neck and is at about the level of the velar
opening. There are 4 large, slightly recurved lips with smooth,
simple edges. There are 4 straight, narrow radial-canals and a
broad circular vessel. The entoderm of the manubrium and ten-
tacle-bulbs is delicate pink and in some specimens a green streak
extends along the outer surface of the entodermal lining of the
radial-canals. This medusa is common at the Tortugas, Florida,
and among the Bahamas throughout the summer. An appar-
ently identical species is found at Suva in the Fiji Islands, South
Pacific, although the Pacific medusa is not so highly colored.

A closely related, if not identical, medusa is described by
Lobianco, 1903, from two specimens drawn from depths of 500 to
600 fathoms near Capri, Bay of Naples, Italy. Dr. Lobianco
kindly permitted me to studythese specimens. The bell is 1 1 mm.
high, 7 mm. wide; thick-walled, with dome-like apex. There are
12 to 13 long tentacles, 4 to 5 times as long as the bell-height.
These have long, hollow, tapering basal bulbs, each with an
abaxial, dark-red ectodermal ocellus. In addition to the long
tentacles there are about 24 to 36 very short rudimentary marginal
tentacles, each with an abaxial ocellus. The 4 radial-canals are
quite wide and with slightly jagged outlines. The manubrium
is half as long as the depth of the bell-cavity. The 4 complexly
folded lips are at the end of a well-developed neck. The outer
surfaces of the adradial gonads are smooth. The entoderm is
strawberry-pink. I can not separate this medusa from Pandea
violacea of Tortugas, Florida, some specimens of which are fully
as pink in color as is the one shown in Lobianco's figure.

FK;. 64. — Pandra vtolacea, after
Lobianco, Mitth. Zool. Sta.
Neapel, 1903.

Pandea maasi.
Tiara, j/)., MAAS, 1904, Result. Camp. Sci. Prince dc Monaco, fasc. 28, p. 13, plate 2, fig. M.

Bell 10 to 13 mm. high and only about half as wide as high, with a smooth exumbrella
surface. Bell-walls quite thick, with an evenly rounded apex, without an apical pro]i-ctmn.
4 thick, hollow tentacles at the bases of the 4 radial-canals. These have large, swollen basal
bulbs, but no ocelli were observed. The tentacles are about as long as the bell-height and

120 MEDUS.E OF THE WOULD.

their shafts are of about uniform width throughout, the outer ends being very blunt, not taper-
ing. There are no other tentacles or marginal appendages. The 4 radial-canals and the ring-
canal are simple, narrow, and straight-edged; without glandular diverticula.

The stomach is wide and barrel-shaped to cylindrical and is about three-fourths as long
as the depth of the bell-cavity. There is no peduncle and the 4 radial-canals run directly into
the stomach without enlarging as they approach its base. The mouth is well developed, but
the lips are simple, without the foldings commonly seen in other Tiarinae.

The gonad is developed as an open network of more or less transverse folds over the ecto-
dermal wall of the stomach, excepting only the neighborhood of the mouth. The gonad is not
distinctly separated into 4 interradial parts, but is more or less fused over the perradial lines,
thus encircling the stomach very much as in medusae of the Codonidae or in Pandca con i en.

The entoderm is yellow, the gonads being deep in color and the tentacles lighter. Other
parts are transparent.

Found by the Prince of Monaco at Bear Island, between Norway and Spitsbergen, in
July, 1898. It is well figured and described by Maas, 1904. Hartlaub, 1907, considers this
medusa to be identical with Sarsia flammea Hartlaub. The general proportions of the two
medusae appear to be much alike, but the details of structure of the gonads and the color of
P. maasi appear to distinguish it from other medusae.

Genus TURRIS Lesson, 1843, sens, amend.

Turr'n+Tiara (in part), LESSON, 1837, Prodrom. Monog. Meduses, Nos. 17, 20 (not published); 1843, Hist. Zooph. Acal.,

pp. 283, 286.

Turris + Oceania (in part), FORBES, 1848, British Naked-eyed Medusae, pp. 21, 26.
Turrh+ Tiara, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 346, 347. — HARTLAUB, 1892, Nachricht. kgl. Gescll.

Wissenschaft. Univ. Gottingen, pp. 20, 22.
Turris, AGASSIZ, A., 1865, North Amer. Acal., p. 164.

Tiara+ Turris+ Catablema, HAECKEL, 1879, Syst. der Medusen, pp. 56, 60, 62.
Turris, FEWKES, 1881, Bull. Mus. Comp. Zoo], at Harvard College, vol. 8, p. 147.
Tiara, WAGNER, 1885, Wirbellosen des Weissen Meeres, p. 78. — MAAS, 1905, Craspedoten Medusen der Siboqa Expedition,

Monoe. 10, p. 14. — BOVERI, 1890, Jena. Zeit. Naturw., Bd. 24, p. 339. (The egg gives rise to 2 polar globules.)
Cu1ablema-\- Turris + Tiara, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 444.
Catablema+ Tiara+ Turris, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, pp. 12, 15, 16.

GENERIC CHARACTERS.

Tiannae with 4 or more marginal tentacles. With 4 interradial horseshoe-shaped gonads
in the stomach-wall. Each horseshoe composed of partially fused swollen ridges.

Lesson, 1843, describes three medusae under the name Turns, only one of which belongs
to the genus as we define it. This one is Turns neglecta of the north Atlantic coasts of Europe.
The name Tiara was also proposed by Lesson, 1843, and is used to describe a medusa which
was first observed by Forskal, 1775, under the name Medusa pileata. It is found off the
Atlantic coast of Europe and in the Mediterranean.

I use Turris as equivalent to Turris Lesson + Tiara Lesson + Catablema Haeckel.
Haeckel, 1879, p. 62, establishes Catablema for a Greenland medusa which he calls Catablema
campanula.

Maas, 1904, who embodies the results of the studies of Vanhbffen, Hartlaub, and himself,
defines Turns as having horseshoe-shaped interradial gonads, the sides of the horseshoe
being made up of partially fused, transverse ridges while the arch connecting the two sides is
composed of an irregular network of ridges. In Tiara the network is lacking, the entire
horseshoe being composed of more or less transverse ridges. The ridges do, however, anas-
tomose to some extent, so that an imperfect, partial network may be formed. Catablema he
distinguishes by the jagged edges of its radial-canals and circular vessel. However, medusa;
of Turns and Tiara often display lagged edges upon their canals, and this character is very
variable and subject to much individual irregularity in development. In the young medusa
of Catablema the ridges forming its interradial horseshoe-shaped gonads tend more or less
longitudinally, but in later development they come to he almost transversely as in Turns
or Tiara. In order to terminate the confusion that has been introduced by these intergrading
and too precise criteria for distinguishing the genera Turris, Tiara, and Catahlrma, I propose
to unite them all under one genus and call it Turns, the name first used by Lesson to dis-
tinguish any of these medusae.

ANTHOMKUUS.E — '1THUIS.
Tubular Synopsis of the Species of Turris.

121

T. papua Lesson.

T.rotunda = Tiara
rotunda Haeckel.*

T.prismatica =
Tiara prismatica
Maas.f

T.reticulata
Haeckel.

T. vesicaria
A. Agassiz.

Size of bell in mm.

28 high, 14 wide.

Pointed apex.

20 high, 20 wide.

Cubical.

20 high, 12 wide.

12 high, 10 wide.

25 high, 12 wide.

Number of tenta-
cles.

8 to 32

8

12 to 1 6

16

16 long, 48 short.

Ectodcrmal, ab-

Present.

Prcbcnt.

?

Present.

Present.

axial ocelli on

tentacle-bulbs.

Character of edges

Serrated.

Smooth.

Serrated.

Smooth.

Jagged.

Character of
gonads.

4 interradial horsc-
shoes of complex,
mainly transverse
swellings which
anastomose to
some extent.

4 pairs of adradial
groups of cross-
folds.

As in T. pilcMta.

8 separate triangular
areas in adradii of
stomach, with
complex anasto-
mosing ridges.

Swollen, transverse,

papilliform,
anastomosing
folds forming net-
work in 8 adradii
of stomach.

Color.

Tentacle-bulbs an.l

Tentacle-bulbE]

?

?

Stomach and ten-

stomach pink to
dark-red. Gonads
pink to dark-
brownish-yellow.

stomach, and
gonads carmine.
Radial-canals and
ring-canal yellow.

tacle-bulbs cin-
namon-brown.
Gonads darker.
<krlli dark-red.

Where found.

Pacific and Indian

Straits, of Gibraltar.

Gulf Stream, At-

South Atlantic nc.u

An tic Ocean to

Oceans in tropical
regions.

lantic.

Tristan
d' Acunha.

southern coast of
New England,
United States.

T.superba Mayer .J

T.coeca Hartlaub.

T. pelagica Agassiz
and Mayer.

T. conifera= Tiara
conifera Haeckel .J

T.campanula =
Catablema cam-
panula Haeckel.

Size of bell in mm.

^ high, 3.5 wide.

^o high, 20 wide.

16 high, 9 wide.

25 high, 2O \v ulr.

20 high, 20 wide.

Number of tenta-
cles.

4 long, 12 short.

24 to 30 of various
lengths.

30 short. All of
same length.

24 to 48. All long.

24 to 48* All long.

Ectodermal, ab-
aiial ocelli on

Present.

Not present.

Not present.

_'

Not present.

tentacle-bulbs.

Character of
edges of radial-
canals.

Smooth.

Jagged.

Jagged.

?

Jagged.

Character of

gonads.

4 interradial horse-
shoes of partially
fused, mainly
transverse ridges.

AsinT. superba.

As in T. vesicaria.

Simple transverse
ridges across the
8 adradii of
stomach.

4 crescent-shaped
gonads in the 4

intcrradii of
stomach. Each
composed of
longitudinal
ridges side bv
side. No trans-

verse folds.

Color.

Stomach emerald-
green and pink to
rose. Tentacle-

Stomach pink,
rose-colored,
wine-colored.

All entodermal parts
light port-wine-
colored.

p

Entodermal parts
yellow.

bulbs rose-color. Tentacle-bulbs

Entire gelatinous , yellow.

substance delicate

*

pink.

\VlnTr found.

Tortugas and
Bahamas in sum-

Mediterranean.

Pacific coast of
Lower California.

Arctic Ocean,
Russia, Green-

Arctic Ocean.

mer.

land.

* Probably an abnnrmal or stunted T. fiilrata.
J Probably ulrntical with T. pilcala.

s this an abnormal individual of T. fileala (?)

122

MEDUS.E OF THE WORLD.

According to Wright, 1859 (Edinburgh New Philosoph. Journ., plate 8, fig. i) and
Hincks, 1868 (British Hyd. Zooph., p. 14, plate 3, fig. l), the egg of Turris neglecta develops
into a Tubulanan hydroid. Clavula gossei (see also Allman, 1871, Monog. Tubularian
Hydroids, p. 259; also Metschnilcoff, 1886, Arbeit. Zool. Inst., Wien, Bd. 6, p. 239) found
that the eggs of Tiara pileata develop into a clava-like hydroid polyp. The hydroid polypites
of Turris (Clavula) arise singly from a creeping stolon. Each polypite is slender, elongated,
and claviform with scattered, elongate tentacles. The coenosarc is invested by a polypary.
It is possible that the hydroid described by Gegenbaur, 1854 (Verhandl. phys.-med. Gesell.
Wiirzburg, Bd. 4, p. 165, taf. I, figs. 3, 4) may prove to be the hydroid of Turris. The short,
clavate polypites arise separately from a somewhat branching stolon, and each has 5 to 8
scattered, slender tentacles. The medusa-buds arise from the stolon and when set free each
medusa has 2 long and 2 immature tentacles, and a line of nematocysts extends up from each
tentacle-bulb to the top of the bell. Gegenbaur calls this form Clcodoru tricuspiJata or
Syncoryna cleodorir. Allman, 1871 (Monog. Tubularian Hydroids. p. 260) calls it Cam-
paniclava.

It is probable that certain species of Perigonimus give rise to medusae of the genus Tun-is.

Corydendrium minor Nutting, 1905 (U. S. Fish Commission Bulletin for 1903, p. 941,
plate 2, fig. I ; plate 7, figs. 8, 9) from the Hawaiian Islands may be the hydroid of some Turris.

Turris neglecta Lesson.

Kfnnin beroe, SLABBER, 1778, Physikal. Belustig., Niirnberg, p. lio, plate 13, fig. 3.

Turris neg/ecla, LESSON, 1843, Hist. Zooph. Acal., p. 284. — FORBES, 1848, British Naked-eyed Medusa1, p. 23, plate 3, figs.

za-2.1.

Oceania globulosa (young?), FORBES, 1848, British Naked-eyed Medusa1, p. 29, plate 3, fig. 3.
Turris neglecta (hydroid ?), GOSSE, 1853, Nat. Rambles Devonshire Coast, p. 348, plate 13, figs. 6-10.
Clavula gossei, WRIGHT, 1859, Edinburgh New Phil. Journ, plate 8, fig. l.
Turris neglecta, HINCKS, 1868, Hist. British Hydroid Zooph., p. 13, plate 3, fig. I. — ALLMAN, 1872, Monog. Tubularian Hydroids,

p. 259. — HAECKEL, 1879, Syst. der Medusen, p. 62.— BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 488, Ibid., 1905,

tome 13, p. 152 (all literature to 1850).

Tabular Synopsis of the Species of Turns — Continued.

T.eurystoma = Cat-
ablema eurystoma
Hacckel.

T. pilrata = Tiara
pileata L. Agas-
siz.*

T. neglecta Lesson.-]

T. digitalis Forbes.

T.brevicornisMur-
bach and Shearer.

Size of bell in mm.

40 high, 20 wj e.

40 high, 25 wide.

6 high, 4 wide.

40 high, 20 wide.

45ni'gh<35 wide.

Number of tenta-
cles.

24 to 48. All long
and 24 to 32 ten-
tacle-bulbs.

12, to 48. Usually
241032. All well
developed.

60 to 70 short.

50 to 100 short.

140 +

Ectodermal, ab-
axial ocelli on

Not present.

Present.

Present.

?

Not present.

tentacle-bulbs.

Character of edges
of radial-canals.

Jagged-

Slightly notched
and jagged.

Jagged.

Jagged.

Character of
gonads.

Developed in longi-
tudinal curtain-
like folds of in-

4 interradial horse-
shoes of mainly
transverse, some-

As in T. pileata.

8 adradia rows of
transverse ridges.

As in T. pileata.

terradial sides of

what anastomos-

short stomach.

ing ridges.

Color.

Stomach and
gonads cherry-
red. Canals and
tentacles bright-
red.

Variable. Stomach
almost colorless,
yellow, red, purple,
brown. Radial-
canals colorless,

Stomach, gonads,
tentacle-bulbs
scarlet, purple.

Stomach purple,
reddish-brown.
Tentacle-bulbs
yellow, orange.

Tentacles, gonads,
and stomach dark-
red, purple.

yellow, green.
Tentacles color-

less, pink.

Where found.

Coast of Greenland.

North Atlantic to

British coasts.

Greenland to North

St. Paul Island,

Mediterranean.

Sea, British

Pribylofflslands.

coasts.

North Pacific.

*T. conifera rotunda, superba and prhmatica are probably identical with T. pileata. Hydroid: Perigonimus ref>ens?
•(•Hydroid: Clavula gossei Wright.

PLATE 13.

Fig. i. Protiara formosn, mature medusa. Tortugas, Florida, May 21, 1907.

Oral view of female.
Fig. 2. Protiara forniosa. One of the marginal cirri showing the chordate

entodermal cells.
Figs. 3 to 5. Podocoryiif fulgnraiis. Agassiz Laboratory, Newport, Rhode

Island, September, 1892.
Fig. 6. Turns pileata, young medusa. Agassiz Laboratory, Newport,

Rhode Island, September 25, 1896.
Fig. 7. Turris vesicaria, mature medusa. Eastport, Maine, August, 1897.

Drawn from life, by the author.

PLATE 13

.

ANTHOMEDU8/E— TDRRI8.

123

Bell 4 to 6 mm. high, 5 to 4 mm. wide. Globular or with a conical apex and bulging
sides; margin somewhat constricted. 60 to 70 short, crowded tentacles, shorter than the

bell-height, with abaxial ocelli. Stomach wide, cruciform in
cross-section, and one-half to two-thirds as long as the depth
of the bell-cavity. 4 simple lanceolate lips, not complexly
tolded. Gonads as in T. pilrtitu. Entoderm of stomach
and tentacle-bulbs reddish-brown to rich purple. Found
or!" the British coasts.

The hydroid was obtained by Grosse and Wright, who
reared it from the egg. The planula is bright crimson.
The hydranths are club-shaped and only about 2.5 mm. high.
They arise singly at intervals from a creeping filiform h\dm-
rhiza and have 12 or more scattered, filiform tentacles. The

Fn.. 65. Hvdroid of
' Hincks,
Zoophytes.

rj negfecta, after hydrocaulus and hvdrorhiza are invested by a perisaic.

Hincks, in British Hvdroid .; .

Hydranths bright crimson.

Turris pileata.
Plate n, fig. 4; plate 13, fig. d.
Medusa pileata, FORSKAL, 1775, Descriptions Animal., p. no; 1776, Icones rerum naturalium, plan- 33, fig. I).

ania pileata+ P. Lesueur, PERON ET LESUEUR, 1809, Ann. du Museum d'Hist. Nat., tome 14, p.

nia (Charvbdea) pisifera, OKEN, 1815, Lehrbuch iler Naturges., Tril 3, p. 125.
ania ampitHacea, SARS, 1835, Beskriv og Jagttagelser, p. 22, plate 4, fig. fta-f.
ra f>,ifia/ii+T. sarsii, LESSON, 1843, Hist. Zooph. AcaL, p. 287.

ania turrita+0. octona+O. episcopates, FORBES, 1848, British Naked-eyed Mediis.r, pp. 27, 28, plate 2, figs. 1-3.
ania coccinea, LEI'CKART, 1856, Archiv. fur Naturges., Jahrg. 22, p. 24.
alidium arnpullaceum + Tiara pileata, A<;ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 352, 347.

0

0:

T

Or

Or

Ph

Tiara smara^dina, HAECKEL, 1864, Jena. Zeit. fiir Naturvv., Bd. i, p. 336.

Oceania pileata, SPAT.NOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 21, tav. 3, fig. i, 2.

Tiata pileata, HAECKEL, 1879, Syst. der Medusen, p. 58, taf. 3, fign. 6-8.— HAMANN, 1883, Zeit. fiir wissen. Zoo!., Bd. 38, p. 426,
taf. 23, fign. 16-20 (development of the planula larva).— METSCHNIKOFF, E., 1886, Embryol. Sludien an Medusen, pp.
29, 48, etc., taf. l, fign. 1-17. — HARTLAUB, 1894, Wissen. Meeresuntersuch. Komni. Mrrrr Ku-l, Helgoland, Neue Folge,
Bd. l, p. 189. — FORSKAL= Oceania epiicopalis FORBES; BROWNE, 1896, Irish Naturalist, p. 180; 1895, Proc. and
Trans. Liverpool Biol. Soc., vol. 9, p. 258. — BEDOT, 1901, Revue Suisse de Zoo]., tome 9, p. 487; Ibid., 1905, tome 13,
p. 150 (all literature to 1850). — BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 11. MAAS, 1904, Result. Camp
Sci. Prince de Monaco, fasc. 28, p. 15, plate l, fig. 9. — BROWNE, 1905, Proc. Royal Soc. Edinburgh] vol. 25, p. 760.

(?) Tiara intermedia, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 277 (Falkland Islands, South Atlantic).

Gervonia octona, FLEMINC, 1823, Edinburgh Philosoph. Journ., vol. 8, p. 299.

Oceania octona, FORBES, 1848, British Naked-eved Medusa-, p. 27, plate 2, figs. 3^-3^. The plate is wronglv laln-lt-d *'O. epis-
copal is ."

Oceania turrita (young medusa), FORBES, 1848, British Naked-eyed Medus.e, p. 28, plate 2, figs, la-zc.

Tiara octona, A<;ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.

Oceania coronala, ALLMAN, 1871, Monog. Tubularian Hvdroids, p. 33, fig. 8.

Tiara octona, H^FCKEI., 1879, Svst. der Medusen, p. 57.

Yon Tiara octona, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 13, plate 2, fig. ll.

Bell about 15 to 40 mm. high and 10 to 20 mm. wide. Side walls quite thin, but there is
usually a well-developed, solid, apical projection, although this may lie absent. This pro-
jection may be conical or cylindrical, with a basal constriction, or pineapple-shaped, etc.
There are 12 to 48, usually 24 to 32, tentacles with hollow, laterally compressed, tapering
basal bulbs. These tentacles are longer than the bell-height and there is an abaxial ocellus
on the outer side of the base of each bulb. The velum is narrow and simple.

The 4 radial-canals and ring-canal are wide, flat, and ribbon-like and their edges are often
more or less notched and jagged. As they approach the sides of the stomach the radial-canals
widen out into funnel-like expansions, so that they embrace the upper halves of the sides of
the stomach in the 4 principal radii.

The stomach is wide, balloon-shaped, and 4-sided, and fills the upper one-half to two-
thirds of the bell-cavity. The 4 lips are at the end of a relatively- narrow, short neck and
are complexly folded, crenated and recurved upward.

The gonads are 4 horseshoe-shaped regions on the sides of the stomach, the sides of each
horseshoe being adradial and the apex aboral and interradial. I he horseshoe consists of an
area of more or less transverse folds which, however, anastomose more or less so as to form
a network of ridges. In young medusz the ridges of the gonads are more or less parallel
and transverse, but in later life they anastomose to form an irregular network.

124

OF THE WORLD.

This medusa is very variable in color. The stomach ranges from yellow to red, or from

brownish-red to purple. The radial and circular canals may be colorless, to yellow or green.

The ocelli are red or dark brownish-purple. The tentacles range from colorless to light-purple.

It is widely distributed over the North Atlantic and in the Mediterranean, but it

appears to be much commoner on the European than along the American coast.

The development has been studied by Hamann, 1883, and Metschnikoff, 1886. Seg-
mentation is total and equal and a single-layered, elongate, ciliated larva is found. The
entoderm develops from cells which wander into the central segmentation cavity from the
hinder end of the larva, so that finally a two-layered planula is formed. The planula attaches
itself and the hinder end develops into a hydrorhiza, while the forward end becomes the
polypite. The hydrorhiza becomes invested with a thin pensarc and the polypite develops

a mouth and 2 or 3 long thin tentacles, which arise at the base
of the conical throat-tube. This stage was reached at the
end of 16 days in Metschnikoft's specimens, and apparently
the hydroid is one ot the Clavidae.

This hydroid may possibly be the very abundant and
widely distributed Pengonimits rrpens (fig. 66) found oft the
Atlantic coasts of Europe from Russia southward, in the
Mediterranean, off the Pacific coast of the United States,
and at the Falkland Islands and Terra del Fuego, South

o

America.

The stems of the hydroid branch and are covered with
a gelatinous investment to which foreign bodies become
attached. The polypites are spindle-shaped, with a single
oral circlet of tentacles. The medusa-buds arise singly from
the sides of the stems.

According- to Hartlaub and Browne the medusa is set

P>

free with but 2 tentacles and 2 small tentacle-bulbs. Forbes
described the 4-tentacle stage as Oceania titrnta. the 8-ten-

• o

tacle condition as O. octona, and the medusa with 12 tentacles

as 0. episcopalu.

Some of the recent literature relating to the hydroid P.
repens is here presented, though it must be borne in mind that this has not been proven to
be the hydroid of the medusa in question.

Pengonimits re pens + P. minutus, ALLMAN, 1872, Monog. Tubul. Hydroids, p. 521, plate II.

Perigommus repens, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 350. (At Trieste, Adri-
atic. Rare. Medusae produced in May). — DU PLESSIS, 1888, Recueil Zool. Suisse, tome 4,
p. 539. — BROWNE, 1896, Proc. Zool. Soc. London, p. 462 (P. minutus ALLMAN = P. repens). —
BONNEVIE, 1899, The Norwegian North Atlantic Expedition, 1876-1878, vol. 26, Hydroids,
p. 41 (from Norway). — CALKINS, 1899, Proc. Boston Soc. Nat. Hist., vol. 28, p. 539, plate
I, figs. 3-3 D (from Puget Sound). — TORREY, 1902, California Univ. Pub. Zool., vol. I,
p. 29 (Pacific coast, California to Washington). — HARTLAUB, 1905, Zoolog. Jahrbiichern,
Suppl. 6, p. 530, fign. K, L, M (from the Falkland Islands and Terra del Fuego).

Turris rotunda.

l>i,in,ra rotunda, QUOY ct GAIMARD, 1827, Annal. des Sci. Nat., tome 10, p. 181, plate 6 A, figs, i, z.

Tiara rotunda, HAECKEL, 1879, Syst. der Medusen, p. 57, taf. 3, fign. 9, lo. — BEDOT, 1905, Revue Suisse de Zool., tome I},
p. 151 (all literature to 1850).

Bell cubical without a pointed apex; 20 mm. high, 20 mm. wide. Bell-cavity only two-
thirds as deep as the bell-height. 8 tentacles much longer than the bell-height, with conical
basal bulbs. A large, dark-red ocellus on the outer side of each tentacle-bulb. Stomach
small, swollen, globular, and confined to the upper third of the bell-cavity. The lips are short,
much folded, and with a row of nettling warts around their edges. The gonads consist of
4 pairs ot longitudinal, swollen areas, each thrown into 8 to 10 cross-foldings, on the adradial
sides ot the stomach. The stomach, gonads, and tentacle-bulbs are carmine, lips rose-red.

1 i'.. i<>>. — Perigonimus repent, after Him k
in British Hydroid Zoophytes.

.\\TIHtMKIirs.K TI'KlilS. I!'.".

Canals yellow. Found by Ouoy and Gaimard, and later by Haeckel, in the Straits of Gib-
raltar. Development unknown.

Turris papua Lesson.

Tuna papua, LFSSON, 1843, Hist. Zooph. Acalephes, p. 283; Prodromus, 1837, No. 36. — EYDOI x I:T SOILEVKT, 1X41.
Voyage de la Bonne, tome 2, p. (t\q, plate i, figs. 1-3. — HAECKKL, 1879, Sysl. der Meduscn, p. 58. — MAAS, 1905, ( i
doten M.-dusen der Xibaga Expedition, Monog. 10, p. 14, taf. l, fig. 13; 1906, Kevue Smsse ilr Zool., tome 14, p. 8X.
BF-UOT, 1905, Revue Suisse de Zool., tome 13, p. 150 (all literature to 1850).

.fyuorea tniiTti, LESSON, 1829, Voyage de la Coqiiille, Zool., p. 127, plate 14, fig. 4.

(?) Tiara ocranica, AGASSIZ, A., and MAYER, 1902, Mem. Mus. Comp. Zool. at Harvard College, vol. 16, p. 141, plate I, fig. I.

(?)Tf<vd intermedia, BROWNE, 1902, Annals and Mag. Nat. Hist., MT. 7, vol. 9, p. 277.

Bell 28 mm. high, 14 mm. wide, with a solid apical projection and thin vertical walls. The
longest tentacles are about as long as the hell-height and have elongate, conical, basal bulbs,
with dark-red ectodermal ocelli on their outer sides. According to Lesson and to Eydoux et
Souleyet there are 8 tentacles: 4 radial, 4 interradial. Maas found 16 tentacles: 4 radial, 4
interradial, and 8 adradial. Agassiz and Mayer tound 32 tentacles: 8 long radial and
interradial, 8 short adradial, and lu very short intermediate tentacles. These differences may
he local variations, but can hardly be due to the medusae being in various stages of develop-
ment, for Agassiz's specimen was only 5 mm. high and had 52 tentacles, while Browne's
and Maas's specimens were 7 mm. high with only 16 tentacles. Moreover, Lesson's medusae
were larger and had only 8 tentacles. The 4 radial-canals are quite wide and have serrau-il
edges. The stomach is wide, swollen, and the lips complexly folded, and about at the level of"
velar opening. The gonads are in 4 interradial, horseshoe-shaped series of transverse swell-
ings on the sides ot the stomach, the open ends ot the horseshoe being directed downward
toward the mouth and the apex being near the aboral end ot the stomach.

The entoderm of tentacle-bulbs and stomach is pink to dark-red. The gonads are pink t<>
dark brownish-yellow.

Widely distributed over the Indian Ocean and tropical Pacific.

It is probable that this form displays considerable variability in the number of its tentacles
and in the color of its gonads. It is closely allied to 'Turns superb a ot Tortugas, Florida, and
mav be identical with T. intermedia Browne, from the Falkland Islands.

Turris prismatica.
Tiara pTiimalifa, MAAS, 1893, Ergeh. der Plankton Exped., Bd. 2, K. c.. p. f»8, taf. 6, fign. 10, 1 1.

Bell about 20 mm. in height, 12 mm. in diameter; 4-sided and prismatic in shape. There
is no apical projection and the bell-walls are remarkably thick. In the single specimen
described by Maas there are 12 tentacles, one at the foot of each of the 4 radial-canals. 3
others in one quadrant, 3 in another, and one in each of the other quadrants. It seems probable
that the specimen is abnormal and that there are normally in tentacles. The tentacles arise at
a slight distance above the bell-margin. Ocelli ( ?) The gonads are similar to those ot Tuna
pilnitii. In the preserved specimen the bell has a 4-sided prismatic shape. Color ( ?) A
single individual of this species was found by the Plankton Expedition on August 4, 1 880., in the
Gulf Stream. May it not be an abnormal specimen of Turris pileata ( ?)

Turris reticulata Haeckel.

Tiara retifitlala, HATI-WEI., 1879, Syst. der Medusen, p. 60, taf. 3, fig. 1 1 .

Medusa bell-shaped, 12 mm. high and 10 mm. broad. There is a small, conical, apical
projection, about 1.25 times as high as it is broad. The stomach is 4-sided and pyramidal,
widest below, and its radial edges are bound throughout their length by the 4 radial-canals.
The 4 lips are large and much folded. There are 16 tentacles which are longer than the width
of the bell, and have large, laterally compressed bases, with ocelli on their outer sides. Their
are 8 separate, triangular gonads in the outer wall of the stomach. The surfaces ot the gonads
display complex anastomosing folds and swellings.

This species was described by Haeckel from preserved specimens obtained in the South
Atlantic near the Island ot Tristan d'Acunha.

12f) MEDUSA OF THE WOULD.

Turris vesicaria A. Agassiz.

Plate ll, figs. 2 and 3; plate 13, fig. 7.

Turris vesicaria, AC.ASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 97; 1865, North Amer. Acal., p. 164, figs. 261-268.—

NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 375, fig. 86 (abnormal medusa).

C.nlablema vesicarium, HAECKF.L, 1879, Syst.der Medusen, p. 64. — GRONBERG, 1898,200!. Jahrb., Abth. Syst., Bd. n, p. 461.
Turris efitcopalit, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 147, plate 3, figs. 1-6.
Turris vesicaria, and T. efiscofalis, HARC.ITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 36, 2 figs.
Catahlema I'tsicarium (young medusa), MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 12, planches I, 2.

Adult medusa (plate 13, fig. 7). — Bell about 25 mm. in height and 12 mm. in diameter.
There is a large, solid, apical projection which varies greatly in shape, being globular in some
individuals, while in others it is cylindrical or conical. Its upper end is often seen to terminate
in a small button-shaped projection. The side walls of the bell are quite thin and flexible.
There are 16 well-developed tentacles, the basal bulbs of which are quite large and are
flattened in a radial direction, so that the tentacle appears to arise from the side of the bell
slightly above the bell-margin. These well-developed tentacles are very flexible, and when
ejctended are about 2 to 4 times as long as the bell-height. These basal bulbs are hollow and
bear each an abaxial ocellus. In addition to these long tentacles there are 48 very short, rudi-
mentary tentacles, which arise from the bell-margin at a slightly lower level than the long ten-
tacles. There are 3 of these rudimentary tentacles between each successive pair of long
tentacles. There is a single dark-red ocellus upon the outer side of each and every tentacle-
bulb, 64 in all. The velum is narrow. There are 4 broad, flat radial-tubes and a broad,
circular canal, all of which display jagged edges. The 4 radial-canals enter the stomach by 4
wide, funnel-like openings. The manubrium is large and urn-shaped and there is no peduncle.
The mouth is surrounded by 4 prominent, crenulated lips. The gonads occupy 4 interradially
situated, longitudinal regions upon the walls of the stomach. Each gonad consists ot a pair
of swollen, papilliform, adradial, ectodermal ridges, the outer surfaces of which are thrown
into complex, but mainly transverse, folds and corrugations. The color of the manubrium and
tentacle-bulbs is cinnamon-brown, the gonads being of a somewhat darker shade. The
ocelli upon the tentacle-bulbs are dark-red.

y otnig medusa (plate 12, fig. 2). — In the youngest medusa observed the bell is about 2 mm.
in height. The walls are thin, and there is a short, conical, apical projection. There are only
4 well-developed tentacles, one at the base of each of the radial-canals. These tentacles have
large conical basal bulbs which are hollow. The axes of the tentacle-bulbs are at a slight dis-
tance above the bell-margin. In addition to the 4 well-developed, radial tentacles, there are 4
intermediate interradial tentacles which arise from the bell-margin at a slightly lower level than
the radial tentacles. The 4 radial-canals are broad and their edges are jagged. The manubrium
is large and quadratic in cross-section. The lips are simple.

In a medusa 3 mm. in height (plate 12, fig. 3), there were 8 well-developed and 8 immature
tentacles. The apical projection was large and miter-shaped, and the lips had folded, notched
margins.

This medusa is probably an Arctic species. It has been found but rarely south of Cape
Cod, and then only in May and June. It is abundant, however, in the harbor of Eastport,
Maine, in August, and ample opportunities for observing its growth were afforded.

The medusa is remarkable for its voracity and will devour Nanomia cara with great
avidity. It is a common thing to observe the medusa in the act of devouring one of these large
Siphonophores which may be many times its own size. Maas, 1904, records this medusa off"
Bear Island, between Norway and Spitzbergen, so that its distribution is probably circum-
polar. Grcinberg records it as being rare in August off" the coasts of Greenland and Spitz-
bergen.

Turris superba (a variety of T. pileata).

Plate 27, fig. 8; plate 28, figs. 3 and 4.

Tiara superba, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 34, plate 16, fig. 39; Memoirs Nat.
Sci. Museum Brooklyn Inst. Arts and Sci., vol. I, No. I, p. 8, plate 2, fig. 1 1.

Bell 5 to 7 mm. high and with a well-developed apical projection. There are 4 long,
hollow and 12 small, solid, rudimentary tentacles. The long tentacles are slender, with taper-
ing basal bulbs. There is an ectodermal ocellus upon the outer side of the tentacle-bulb of

ANTHOMEDU8.*— TURRI8.

127

each of the 1 6 tentacles. The velum is well developed. There are 4 broad, flat, smooth-edged
radial-canals and a broad, simple, circular vessel. The 4 radial-canals enter the stomach by
4 wide tunnel-like openings. The manubrium is wide and tills the greater part of the bell-
cavity. The 4 lips are recurved and their edges much tolded. The 4 gonads are developed
on the interradial sides of the stomach. Each gonad is horseshoe-shaped, the apex of the
horseshoe being uppermost, and the sides composed of partially fused, transverse ridges upon
the ectoderm of the stomach-wall. The entire gelatinous substance of the medusa is of a
delicate rose-pink.

The entoderm of the manubrium and tentacles is of a rich rose-color and the entodermal
nut' of the stomach is emerald-green. This medusa is quite common at Tortugas, Honda,
and among the Bahamas in the summer months. It is one of the most beautiful of American
medusae.

Turris breviconis Murbach and Shearer.

Turrit hrei-iconis, Mi'RBAfl! AND SHKAKKK, 1902, Annals ami Mag. Nat. Hist., ser. 7, vol. 9, p. 73; 1903, Proc. /..nl. s.u .
London, vol. 2, p. 170, plate 18, figs. I, 2.

Bell cubical, 45 mm. high, 35 mm. wide. More than 140 tentacles in double rows, with
contractile muscles on their inner sides and enlarged ectoderm on their outer sides. No
ocelli in preserved specimens. Gonads and stomach occupy less than halt of the upper part
of the hell-cavity. 4 broad radial-canals with unbranched lateral diverticula. Bell bluish,
tentacles, gonads, and stomach dark-red or purple.

St. Paul Island, Piihylott' Islands, North Pacific.

Turris coeca Hartlaub.

Turns rrecu, HARTIM-R, 1892, Nachrichten kgl. Gesell. Wissen. Gimingen, p. 19, fig. i.— MAAS, 1904, Result. Camp. Sci.

Prince de Monaco, fasc. 28, p. 17, planchr I, fig. 5.
Tinrclhi fHirlhfrifjf'tifti, TRINCJ, 1907, Archiv. Ital. Anat. Embr. Firenze, vol. 5, p. 533 (development and structure of ooevtes).

Bell about 30 mm. high and with 24 to 30 tentacles of various lengths. The tentacle-
bulbs l<ifk ocelli. The radial-canals are wide and flat and there
are a few globular lobes along their edges. The stomach is
wide and the lips complexly folded. The 4 interradial gonads
are composed of horseshoe-shaped swellings. The ridges are
lateral cross-folds in the 8 adradii, but in the mterradius at the
apex of the horseshoe, near the base of the stomach, they form
a reticulated network.

The stomach is rose-colored to wine-colored, and the ten-
tacle-bulbs are yellow.

Found at Naples and Monaco, Mediterranean, in Febru-
ary and March.

Turris pelagica Agassiz and Mayer.

Turrit prla^irn, AGASSI?, A., and MAYF.R, 1902, Mem. Museum Comp. 7.ool. at
Harvard College, vol. 2(1, p. 142, plate \, fig. 2.

Bell id mm. in height and o mm. wide, and the sides are

O *

I'K,. (17. --Gonail of Tmrii creca, after barrel-shaped, being wider at the middle than at either end.
Maas, in Result. Camp. There is a small, solid, apical projection. The bell-walls are
very thin and <|uite flexible. There are about 30 short tentacles,

less than half as long as the bell-height, all being of the same size. These tentacles all arise
from the bell-margin and their bases are large and conical. There are no ocelli. The velum
is well developed. There are 4 radial tubes which are flat and qujte broad, being narrower
near the circular canal than at any other place. Their outer edges are jagged, excepting in
the narrow parts near the circular canal. The manubrium is large and fills the greater
part of the bell-cavity. It is joined to the 4 radial-canals by means of 4 wide funnel-shaped
ducts. The gonads occupy 8 adradially situated rows which extend about two-thirds of the
distance from the inner apex of the bell-cavity to the velar opening. The outer surfaces
of the gonads are transversely folded and give rise to numerous papillae. There are 4 well-

12S MF.nrs.r, OF THF, WORLD.

developed, complexly folded lips. The lips and tentacle-bulbs are of a light port-wine color
.nul the gonads and radial-canals are of a still lighter shade. A single specimen of this
medusa was found in a surface haul on August 26, 1899, off the Pacific coast of Lower Cali-
fornia; N. lat. 51° 10', \V. long. 125°. It is separated from T. papua and T. [>ilf<ita by the
absence of ocelli.

Turris conifera Haeckel.

Tiara coni frni, HAF.CKFI., 1879, Syst. der Meilusen, p. 59.— FEWKES, 1888, Report on Medusa? of the Lady Franklin Bay Ei-
pi-dition, p. 43.— LFVINSF.N, 1893, Vid. Meddel. Nat. For. Kjobenliavn (5), Bd. 4, p. i44.--GKijNiir.RU, 1898, Zoolog.
[ahrh., Abth. Svst., Bil. i I, pp. 454, 460. — LINK", 1904, Zool. Anzriger, Bd. 28, p. 215.

Bell cone-shaped, with a large, conical, apical projection. It is about 25 mm. in height
and 20 mm. in diameter. There are 24 to 48 tentacles which are longer than the bell-diameter
and which have wide, conical bases. Ocelli ( ?) The manubrium is quadratic in cross-
section and is somewhat longer than wide. The base is wide and cruciform, and the 4 radial
coiners are wide where the radial-canals enter the stomach. The 4 lips are very prominent
and are thrown into complex crenulations and folds. The gonads are composed ot about
10 pairs of regularly arranged, simple transverse or oblique ridges across each ot the 8 adradial
regions of the stomach and are separated in the 4 interradn.

This form is very closely related to Turris pileata, with which it may, indeed, prove
identical. Haeckel says that it is distinguished from both T. pileata and T. reticulata by the
regular feathered gonads and by the smallness of the 4 funnel-like origins of the radial-
canals where they join the stomach. The oral lappets are smaller and less crenulated than
in 7 . pili-titii or T . rrticulatn. The conical, apical projection is half as high as the bell. Color ( ?)

This species is found off the coast of Greenland. Linko, 1904, also found it in Barents
Sea, north of Lapland, and its distribution is, therefore, probably circumpolar in common
with other Arctic species ot Turns.

Turris campanula.

Calablema campanula^ HAF.CKFJ, 1879, Syst. der Medusen, p. 63, taf. 4, fign. 4, 5. — LF.VINSEN, 1893, Vid. Meddel. Nat. For.
Kjohenhavn (5), Bd. 4, p. 144. — GRONBERG, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 1 1, p. 460.— LINKO, 1904, Zool. An-
zeiger, Bd. 28, p. 21?.— BF.DOT, 1901, Revue Suisse de Zool., tome 9, p. 482; Ibid., 1905, tome 13, p. 132 (list of refer-
ences to 1850).

Bell about 20 mm. wide and 20 mm. in height. There is a well-developed, conical,
apical projection, which is hollow and contains a prolongation ot the gastric cavity. There
ate 24 to 48 tentacles which are longer than the bell-diameter. Their bases are long and
thick. No ocelli. The 4 radial-canals and the circular tube are wide and flat and have
numerous complex, or simple, blindly-ending side branches resembling those of T. cury-
stomn. The manubrium is wide and shallow and the 4 lips are thrown into complex
crenulations and folds. The 4 gonads are found within 4 crescent-shaped swellings upon
the 4 interradial sides of the stomach. A number of simple, straight, longitudinal folds or ridges
extend across the crescent-shaped gonad; the horns of each crescent are directed downward,
toward the velar opening. The manubrium, gonads, canals, and tentacles are yellow.

This species is found off the coast of Greenland and at Spitzbergen, and Linko found
it in Barents Sea, north of Lapland.

Turris eurystoma.

Calablema eurystoma, HAECKEL, 1879, Syst. der Medusen, p. 64, taf. 4, fign. 6, 7. — LF.VINSEN, 1893, Vid. Meddel. Nat. For.
Kjobenhavn (5), Bd. 4, p. 144. — GRONBERO, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 11, p. 462.

Bell about 20 to 25 mm. in diameter, and spheroidal, being flatter than it is high. There
is a spheroidal, apical projection which varies in size from about one-fourth to almost as large
as the bell itself. This apical projection is always solid. The sides of the bell are quite thin.
There are about 24 to 48 well-developed tentacles which are much longer than the bell-
diameter. These tentacles have long, conical basal bulbs. In addition to the long tentacles
there are usually 24 to 32 small, immature, or rudimentary tentacle-bulbs which alternate
with the long tentacles. Ocelli ( ?) There are 4 radial-canals, each of which is about 3 mm.
wide in the middle and only 2 mm. wide at either end. Both the radial-canals and the circular

AVTHHMKWS.TC — TTTJIMS.

129

tube give rise to numerous simple or branched, blindly-ending side branches ot o.} to i mm.
in length. Some ot these side branches are simple sacs, but others give rise to secondary
branches. The stomach is wide and cruciform, but the side walls are remarkably short, so
that the mouth is a gaping, cruciform slit, reminding one of" the mouth-opening of Stauro-
pliwa. The 4 interradial sides of the stomach form 4 curtains within which the sperm or
ova are developed. The tree, lower edges of these curtains are crenulated and thrown into
complex folds. The stomach and gonads are cherry-red and the canals and tentacles are
light-red in color. This species is found on the coast of Greenland.

Turris digitalis Forbes.

Ttirris <//t;fM/is, FORHF.S, 1848, British N.ikei!-eyed Medusa-, p. 21, plate 3, figs. lil-lt. — A<;ASSIZ, L., 1X62, Cunt. Nat. Hi t
U. S., vol. 4, p. }4<>. H\KI k M , 1*74, Sv\t . .Icr Mr ilusen, p. f»l , taf. 4, figs. 2, 3. — \>* VINSI N, lS»n, Vui . Mrddrl . N .M
For. Kjdhenh.ivn (5), Bd. 4, p. 144. -Km KIMUKC., iSSo, Vergleu hen, I phvsiolog. Suidien /u Turnv, etc., Aht. 5, p. 124
(reactions to curare and strychnine).

Bell cylindrical, with a well-developed, conical, apical projection. It is 30 to 40 mm. in
height and 15 to 20 mm. in diameter. There are 50 to 100 short tentacles, which appear as if
arranged in two rows owing to the tact that the bases ot the larger ones extend a slight dis-
tance up the sides ot the bell above the margin.

KM.. 68. — Turrii digitalis, after Haeckel, 1879.

Fn;. 6<). — Corri* i\ilf,/>liilnilr>iat after Haeckel, in S\st. »lrr MeJuseti.

There are 8 strands ot longitudinal muscle fibers within the wall of the subumbrella.
These strands lie close to, and upon both sides ot, the 4 radial-canals. The radial-canals are
broad and flat. In its upper halt each radial-canal is a simple, flat tube with a straight edge.
In their lower halves, however, the canals become broader and give rise to many small side
branches which end blindly. The manubnum is very large and fills the greater part ot the
hell-cavity. The mouth extends to the level of" the velar opening. There are 4 prominent
crenulated lips. The gonads occupy 4 interradially situated, longitudinal regions upon the
wall of" the stomach. Each gonad consists ot" a double row of papilliform ridges, the outer
surfaces of which are thrown into complex folds and corrugations.

The stomach is purple or reddish-brown and the lips are rose-colored. The gonads
are darker in hue than the stomach. The tentacle-bulbs are orange or yellow and their
outer ends are white. The bell is transparent. The 8 muscle-strands in the subumbrella
are yellow.

130 MEDUS.TC OF THE WORLD.

The species is found in the North Atlantic. It has been taken off the Shetland Islands,
between Greenland and Iceland, between Iceland and the Hebrides, in the North Sea, and off
the west coast of Greenland. It is probably an Arctic form, of circumpolar distribution.

Genus CONIS Brandt, 1834.

(.'own, BRANDT, 1834, Recueil des Actes-seances publ., Acad. Imp. Sci. St. Petersbourg, p. 19 (of the "separate"); 1838, Mem.
Acad. Imp. Sci. St. Petershourg, ser. 6, vul. 4, par. 2, p. 355. — HAF.CKKL, 1879, Syst. der Medusen, p. 55.

The type species is Conis niitrnta Brandt of the Bonin Islands, North Pacific.

GENERIC CHARACTERS.

Similar to Pandea, but the tentacle-bulbs give rise to special ocellar clubs, which project
outward around the margin. The gonads consist of simple, longitudinal swellings on the
sides of the stomach and have smooth outer surfaces. The hydroid is unknown.

Conis mitrata Brandt.

Conit mitrata, BRANDT, 1834, Recueil Actes publique, Acad. Sci. St. Petersbourg, p. 19 (separate); 1838, Mem. Acad. Imp.
Sci. St. Petersbourg, ser. 6, vol. 4, par. 2, p. 355, pi. 2. — HAECKEL, 1879, Syst. der Meduseo, p. 55. — BEDOT, 1905,
Revue Suisse de Zool., tome 13, p. 133 (literature 1834-50).

Bell 50 mm. high, 35 mm. wide, with a short, conical apex. 64 short tentacles, each with
a club-like ocellar bulb projecting from the outer side of the basal bulb at the margin. The
tentacles are shorter than the bell-height. The stomach is barrel-shaped, with 4 not very
complexly folded lips, having their margins studded with nematocyst-warts. Lips are at a
level about half-way between the velar opening, and the inner apex of the bell-cavity. Stomach
light-red. Tentacle-bulbs blue. Ocelli black. Bonin Islands, North Pacific.

Conis cyclophthalma Haeckel.
Cmit r\rlo[>hihalma, HAECKEL, 1879, Syst. der Medusen, p. 55, taf. 4, fig. i.

Bell pyriform, with thin sides and without an apical projection; 15 mm. high, 12 mm.
wide. There are 52 tentacles with very elongate, conical basal bulbs, one-third as long as the
tentacles themselves. The outer ends of the tentacles are thin and flexible while the basal
bulbs are stiff. The tentacles are longer than the bell-height. At the base of each tentacle
on the abaxial side of the margin there is a short club-like projection which contains a black,
ectodermal ocellus. There are 4 straight-edged radial-canals and a simple ring-canal. The
stomach is very large and almost fills the bell-cavity. There are 4 very large, complexly
folded lips with their margins studded with nematocyst-warts. The lips are at about the
level of the velar opening. (See fig. 69.)

Haeckel figures and describes 4 longitudinal, swollen gonads on the "perradial" sides
of the stomach. Their outer surfaces are smooth and they are filled with ova. He represents
them as being completely separated in the interradii. The gonads are probably adradial or
interradial ( ?). Color ( ?)

Haeckel studied a preserved specimen in the Museum of Copenhagen. It was obtained
near Gibraltar in lat. 30° iff N., long. 2° 23' W.

Genus CALYCOPSIS Fewkes, 1882.
Calycopsis, FEvvk'ts, 1882, Bull. Mus. Comp. Zoo], at Harvard College, vol. 9, p. 304.

The type species is Calycopsis typti Fewkes, from the Gulf Stream off the coast of the
United States.

GENERIC CHARACTERS.

Tiarinae with 16 separate, simple, unbranched radial-canals; 4 radial, 4 interradial, and
8 adradial. Gonads are composed of a row of leaf-like, transverse foldings along each of the
8 adradial sides of the stomach. Lips ( ?) Tentacles simple. Ring-canal simple, without
blindly-ending diverticula. Development unknown.

ANTHOMEDUSjE— CALYCOP818.

131

Calycopsis typa Fewkes.

(,Wv<'//m< t\f><i, l''>.wKts, lSS2, Bull. Mus. Comp. Zool. at Harvard College, vol. 9, p. 304, platr i, fig. 54.

Bell higher than a hemisphere, with evenly rounded top and vertical sides. External
surface smooth, bell-walls thin, thicker above than near the margin. Size ( ?) There are 16
radial-canals and 16 marginal tentacles, one at the base of each canal. These tentacles are

about as long as the bell-height and are flexible with some-
what club-shaped outer ends and very small basal bulbs.
No lithocysts or marginal bodies other than the id ten-
tacles. The stomach is very wide and fills almost the whole
ot the inner half of the bell-cavity. The gonads are devel-
oped upon the sides of the stomach and appear as a series
of transverse, leaf-like ndges on both sides of each of the 4
principal radial lines of the stomach. They are thus
adradial in position and their manner of folding resembles
the condition seen in the gonads of Turns. The sides of
the stomach are bound to the subumbrella by 4 radially
situated partitions. There are 16 straight, narrow radial-
canals: 4 of these are perradial, 4 are mterradial, and 8
adradial. In alcoholic specimens the ovaries were dark-
brown. Tentacles and bell-walls white with a tinge of
bluish color.

4 specimens were found by the Allxitross, in 1880 81,
off the Atlantic coast of the United States, in deep water along the Gulf Stream.
Form of lips ( ?) Size ( ?) Ocelli ( ?)

Fi(i. 70.— Cai\rof>si* t\f>ti, after Fewkes, in
Bull. Museum Comp. Zoo), at
Harvard College.

Subfamily MARGELIN^ Haeckel, 1879.

Oceanidx with oral tentacles and with 4 simple, unhranched radial-canals. Gonads
on the mterradial, or adradial sides of the manubnum; the mature genital products are
tound in the ectoderm.

i . Tribe Cvttftdi:

Marginal tentacles arise singly from bell-margin and are not grouped into clusters. Oral tentacles are unbranched, simple, or

degenerate. They are found in all forms excepting in the degenerate medusa; of Stvlactis.
Cyttris, ESCHSCHOLTZ, 1829. Cyt<ris+ Cubogaster HAECKEL. 4 marginal tentacles. Oral tentacles present. Hydroid

unknown.
Podocorvne, SARS, 1846 = Dysmorf>hosa± Cytcrandra HAECKEL, 1879. 8 or more marginal tentacles. Oral tentacles

present. When present the peduncle is solid. Hydroid: Podocorynt.

Turritopsis, McCRADY, 1856. 8 or more marginal tentacles. Entodermal walls of the 4 radial-canals, above stomach,
are composed of highly vacuolated cells forming a peduncle. Mouth studded with a row of nematocyst-bearing
knobs. Hydroid: Dendroclara.
Oceania, sens., KOLLIKER, 1853; GECJENBAUR, 1856. Medusa similar to Turritopsh, but with a simple, solid, gelatinous

(not vacuolated) peduncle. Hydroid: Clava-like.

Stvlactis, ALLMAN, 1864, degenerate medusae with 4 to 8 rudimentary marginal tentacles and no oral tentacles. Medusa:
mature upon liberation from hydroid. Hydroid: Stylactts; closely related to Podocoryne.

2. Tribe Thamnostomeidi:

With branching oral tentacles. With simple marginal tentacles, not grouped in clusters.

Thamnost\ltis, HAECKEL, 1879. 2 diametrically opposed, marginal tentacles. Development unknown.
Thamnitis, HAECKEL, 1879. 4 radially situated, marginal tentacles. Development unknown.

L\mnorea, PERON and LESUEUR, i8og= Ltmnorea+ Thamnostoma, HAECKEL, 1879. 8 or more marginal tentacles.
Development unknown.

3. Tribe BougatnviUidi:

Marginal tentacles grouped into clusters. Each cluster consisting of 2 or more tentacles. The 4 oral tentacles may be simple

or branched.
Bougainvillia, LESSON, 1836= Margflis + Lizusa± Hippocrene, HAECKKL, 1879. With branching, oral tentacles. With 4

radially situated clusters of marginal tentacles, all of one sort. Hydroid: Bougainvillia.
Ntniopsis, L. AGASSIZ, 1849. With branched oral tentacles. With 4 radially placed clusters of marginal tentacles,

consisting of a median pair of club-shaped tentacles and a number of lateral filiform tentacles in each cluster.

Hydroid: Bougainvillia,
Ratfikfti, BRANDT, 1857 = Mnr^fll turner Lizzia+ Lizztlla+ Rathkeu, HAM-KM., 1^79- With X dusters of marginal ten-

tacles. Mature hydroid unknown.
Cfiiarellat MAAS, 1X97. With 16 (8 double) clusters of marginal tentacles. Circular can.il gi\rs off 4 intrrradi.il, blindly

ending, centripetal branches.

132

MKDUS.E OF THE WUHLI).

Genus CYTjEIS Eschscholtz, 1829.

\_uiiiu. /juui . tii iidi v di u \^<_mcgc, vui. ji, u< iwi . uttuvv i> t, i ywjj ixcfi . i \_c^iun i cd

London, Supplementary Report 27, p. 135. — CHUN, 1895, Bibliotheca Zoologica, Heft 19, p. 11 (law of budding).
(1 \tiris. + Cubogasterf HAECKEL, 1879, Syst. der Meduscn, pp. 73, 75.

Aron Cytais, MAVER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 39.
C.\t,ris, VANHOFKEN, 1891, Zool. Anzeiger, Bd. 14, p. 444.
\on C \tj-is, SARS, 183^, Beskriv og Jagttag., p. 28.

Ctibogasttr, HAECKEL, 1864, Jena. Zeit. fur Naturw., Bd. i, p. 341 (vuung of Cy/.r;.0-
\igritina, STEENSTRUF, 1837, Acta et Catal. Musei Hafmensis, Hauniic.

GENERIC CHARACTERS.

MargeliiiEE with 4 simple, radially situated, marginal tentacles and with 4 or more simple,
unbranched oral tentacles. The hydroid is unknown.

The type species is Cyta-is tetrastyla Eschscholtz, 1829, of the Mediterranean and eastern
parts of the tropical Atlantic. Eschscholtz states that it has 8 simple oral tentacles and 4
marginal tentacles, but he figures at least 10 oral tentacles. More recently species of Cyttcis
have been found with 8, 16, or even 32 oral tentacles.

Tubular Synopsis of the Medusae of Cytirts.

C. tetrastyla

C.atlantica =

C. pusilla

C. vulgaris

C. herdmani

Eschscholtz,

C. macrogaster

Gegenbaur, 1856.

Agassiz and May-

Browne, 1905.

1819.*

Haeckel, 1879

er, iSgg.-f-

= C. tetrastyla

Eydoux ct Soule-

yet, 1841

= C. nigritina

Haeckel, 1879^

Shape and size of

Cubical to domelike

Pyrtform. 5 high,

Oval. 4 high, 3

Prismatic, flat above

Oval. 3.5 high, 3.5

bell in mm.

2 to 3 high, 2 to 3

6 wi e.

wide.

5 high, 3.5 wide.

wide.

wide.

Character of the 4

Similar each to each.

As in C. tetrastyla,

2M- long. With

IT long, with large

As in C. atlantica.

marginal tenta-

2r+ long. Basal

but with large

small basal bulbs.

basal bulbs.

cles. Length in

bulbs small.

pynform basal

terms of bell-

swellings on ex-

raJius (r).

umbrella above

each tentacle.

Shape and size of

Flask-shaped,

Very large, flask-

Stomach oval.

Stomach oval to

Wide flask-shaped,

manubrium.

mounted on

shaped. Two-

Mounted upon

spindle-shape ',

on short peduncle

peduncle. Mouth

thirdsto i .5 times

conical peduncle

half to two-thirds

Two-thirds as

at level of velar

as long asdeptli of

not quite as long

aslong asdepth of

long as depth of

opening.

bcll-cavitv. No

as stomach.

bell-cavity.

bell-cavity.

peduncle in

Mouth at middle

Young medusa has

mature medusa.

of bell-cavity.

short peduncle, but

this is not seen m

mature animal.

Number of oral

4 to 6

8 to 32

12 to 24

32

50 to 60

tentacles.

Character of

4 interradial swell-

Numerous medusa-

4 interradial swell-

8 adradial gonads

4 perradial? or 8

gonads and

ings.

buds on upper

ings.

near base, on sides

adradial ? g<>n.i;K

medusa-buds.

half of manubrium.

of stomach.

No medusa-buds.

Color.

Stomach and tenta-

Stomach yellowish.

Stomach and gonads

Stomach and tenta-

?

cle-bulbs brown.

Gonads and tenta-

vellowish-brown.

cle-bulbs red and

cle-bulbs violet-

Tentacle-bulbs

green. Abaxial

brown, red, or

black.

entoderm red,

nearly black.

superficial green.

Where found.

Mediterranean, and

Mediterranean to

Mediterranean.

Fiji Islands to Malav

Coast of Ceylon.

tropical Atlantic.

Atlantic to

Archipelago, trop-

Canary Islands.

Brazilian coast.

ical Pacific.

*Probably young and immature. Possibly young of C. atlantica?

•f For details see text.

AXTHO.MKDI-S.K — cvr.Kis. i:j;;

The gonads ot C\t<cis are developed upon the mterradial sides of the stomach. In
('\tiris vulgaris, according to Maas, 1905, they occur as 4 pairs of slightly swollen, corrugated
regions, a pair being found on each interradial side of the stomach. The mature genital
products are found in the ectoderm ot the stomach-wall. Many of the species develop medusa-
buds upon the interradial sides ot their stomach. Haeckel's statement that the gonads are
radial in position appears to be erroneous.

Cyttris is closely related to PoJocofyne and is distinguished only by its smaller number
(4) of marginal tentacles, whereas PoJoforync has 8 or more marginal tentacles. MOICOMI
there are commonly more than 4 oral tentacles in ('.\ttcis, whereas the majority of the species
ot Podocoryne have only 4 oral tentacles. C\t<ns is possibly the original genus from which
I'tJvi-rji vii,- and more complex forms may have been derived.

Cttbrjgtisti-r Haeckel, 1864, appears to be only the young ot (,'v/<r/>, excepting " (*.ul»n\:i>t, i
gemmascens" which I take to be an abnormal young Rathkc/i octoputittntu with only 2 well-
developed and 6 retarded marginal tentacles. Broch, 1905 (Bergens Museums Aarbog. No.
11, p. 4), states that the marginal tentacle-bulbs may range from 4 to 8, and that well-
developed tentacles may be absent.

Cytaeis tetrastyla Eschscholtz.

( 'v/,M \ trrr<i*t\l<i, Ls( ns( HOI. rz, 1829, Syst. der Acalrphrn, p. 104, taf. 8, fig. 2. — HAM K M , i S^«j. S\ i . ,lcr MrduM-n, p. ?} .

BKDOT, 1905, Revue Smsse dr Zool., tome 13, p. 133 (literature, 1829-50).
Bougainrillca mcditerranea, BUSCH, 1851, Bcobacll. wirbell. Seethiere, p. 21, taf. 2, fig. 10.

This medusa is recorded trom the Mediterranean and the eastern parts ot the tropical
Atlantic. For details see tabular synopsis ot the medusae ot Cyttris.

Cytaeis atlantica.

iVigriiinti (itltintift2> STKKNSTRUP, 1837, Catal. Musei Hafniensis.

C\(.rn ici',i^\l,i, KVDOUX et SOULEYET, 1841, Voyage de la Botnir, tonic 2, p. ^41, pl.ite 2, figs. 4 to i^.

C.M.ris marrogauer, HAKCKEL, 1879, Syst. der Medusen, p. 74, taf. 6, fig. I.— CHIN, 1X1)5, Bihliotheca Zo-ilogna, Mi-ft iij, p. 4^

(budding of medusa from the manubrium).
(;\[,ris nigritina, HAECKEL, 1879, Syst. der Medusen, p. 74, taf. 6, fign. 2-5. — MAAS, 1904, Result. Cam p. Sa. Pririo- il<- Mi'iiani,

l.i i . 28, p. 8, plate I, figs. 3, 4.

Bell bluntly conical or pear-shaped, about 6 mm. high and 5 mm. wide. There are 4
stout radially placed marginal tentacles longer than the bell-diameter. The bases ot these
tentacles are very thick and extend up the sides ot the bell as large, bulbous, pear-shaped
swellings. These swellings bear a large crescent-shaped mass of entodermal pigment on
their lower, outer sides. These bulbous swellings are each about one-eighth to one-fourth as
high as the bell. The velum is well developed. There are 4 simple, straight radial-canals
and a simple ring-canal. The manubrium is large and lacks a peduncle. It is 0.5 to i.^
times as long as the depth of the bell-cavity. Normally it is probably about as long as the
depth of the bell-cavity. There are numerous medusa-buds upon the upper halt ot the stomach-
wall and these are seen in various stages ot development. 1 he mouth is surrounded In .1
circlet of 8 to 32 short, slender, oral tentacles, each of which terminates in a nematocyst-knob.
The stomach is reddish-brown and the tentacles brown, with tentacle-bulbs nearly black.
This medusa is found in the Mediterranean and Atlantic trom the shores ot Africa to flu-
Brazilian coast.

Haeckel describes 4 gonads at the points ot |uncture ot the 4 radial-canals with the
stomach, but this is probably erroneous, tor it is improbable that the gonads are m the principal
radii.

Haeckel distinguishes C. nigritimi with manubrium half as long as the depth ot the
bell-cavity, and with 16 to }2 oral tentacles; and ('.. niiu-rngantfr with manubrium 1.5 tinns
as long as the depth of the bell-cavity and with 8 to 16 oral tentacles. _ In 1904, Maas dcsciilu s
a very similar medusa, under the name ot (',. nigi'itnm, in which the manubrium is about
two-thirds as long as the depth of the bell-cavity and has i i oral tentacles. These medusa-
are so closely similar in all other respects that I am of the opinion that they are men-l\
growth and contraction phases of one and the same species. Following the usual rule of pre-
cedence, it appears that the specific name should be "atlantica," this being the designation
proposed by Steenstrup, 1837.

134

MEDUS.E OF THE WOULD.

Cytaeis pusilla Gegenbaur.

(.'v/.n'j fuiilla, Gr.iiiNBAi-R, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 228, taf. 8, fig. 8. — KEKERSTEIN UNO EIILERS, 1861, Zoolug.

Beitrag., p. 84, taf. 13, fign. 8, 9. — HAECKEL, 1879, Syst. der Medusen, p. 75.
(?) Cubogasler diuonema (young medusa), HAECKEL, Ibid., p. 76.

This medusa is recorded from the Mediterranean. For details see tabular synopsis of
the medusae of Cyttcis.

Haeckel describes "Cubogaster dissonema" a small medusa 2 mm. high, with 2 well-
developed and 2 immature or rudimentary marginal tentacles, and 8 oral tentacles. This
he found at Croisic, Bretagne, Atlantic coast of France. It may be the young of C. pusilla ( ?)

7'-

Flc. 71. — Cvttfis atlantica, after Maas, in Result. Camp. Sci. Prince de Monaco.

FIG. 72. — Cytefis " macrogaster" ((7. atlantica), after Haeckel, 1879.

FIG. 73. — C.ytas'n pusilla, after Gegenbaur, in Zeit. fur wissen. Zool., Bd. 8.

Cytseis vulgaris Agassiz and Mayer.

Cyttsis vulgaris, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161, plate 2, figs. 3-5. — MAAS,
1905, Craspedoten Medusen der Siboga Exped., Monog. 10, p. 8, taf. i, fign. 4-7; 1906, Revue Suisse de Zool., tome 4,
p. 85, fig. 3, plate 2.

Mature specimens of this medusa are described by Maas from the Malay Archipelago.
Agassiz and Mayer described only half-grown medusae from the Fiji Islands.

Mature medusa. — Bell 5 mm. high, prismatic, 4-sided, and with a flat top. There are
4 thick, radially placed, marginal tentacles, each somewhat shorter than the bell-height. These
tentacles end in blunt tips and have large hollow basal bulbs. Maas describes a thick lens-
shaped swelling on the abaxial side of each tentacle-bulb, but his specimens are somewhat
contracted through preservation in formalin. The basal bulbs of the living medusas studied
by Agassiz and Mayer were large and swollen, but lacked the lenticular, ectodermal swellings.
On the other hand Agassiz and Mayer found only half-grown medusae, and it is possible that
the peculiar form of the basal bulbs described by Maas may be characteristic of the full-grown,

ANTHOMKDI s.K (VI.KIS, I'l i|« M •( >IM NK. 135

but not of the half-grown, medusa. The velum is well developed. The 4 radial-canals and
ring-canal are slender and simple. The manubrium is spindle-shaped and shorter than the
depth of the bell-cavity. In the halt-grown medusa there is a short conical peduncle, but this
was not seen by Maas in his large specimens.

The mouth is at the extremity of a bluntly pointed, conical neck, and is a simple round
opening. 32 or more short, slender, filiform, unbranched oral tentacles arise from a zone
above the mouth. Each oral tentacle is solid and ends in a knob-like cluster of ncmatocysts.

1 he gonads are found on the upper interradial sides of the stomach and appear as a pair
of longitudinal swellings in each of the 4 interradii. They are not developed on the 4 radial
lines of the stomach. The eggs protrude over the surfaces.

In the formalin specimens studied by Maas, the gonads were yellow, the axial entoderm
of the stomach dark-red, as were also the entodermal parts of the tentacle-bulbs. There is
some green pigment in the ectoderm of the tentacle-bulbs. In the immature specimens from
Fiji, studied from life by Dr. Agassiz and his assistant, the entoderm of the outer part of the
manubrium and of the tentacle-tips was green, while there was red pigment in the axial part of
the stomach and in the tentacle-bulbs and adjacent parts of the 4 radial-canals. In the young
specimens from Fiji the oral tentacles ranged from 4 to id, while in those found by Maas in
the Malay Archipelago, they ranged from 8 to 32.

Found in abundance on the surface among the Fiji Islands in November to January.
Maas found numerous specimens in the Siboga collection from the Malay Archipelago,
Sulu, Damar, Daiam. Sabuida, Gisser, Mampa Straits, and Saleyer; and from Amboma, 1906.

Cytaeis herdmani Browne.

(.'w//\ ftciil'tifini, BKDWNK, 1905, IV.ul OvMer I'ishenes, GuH nf Mjnaar, Km . Soi . LMIU|IUI, Suppl. Report 2^, p. i;^, pl.itr I.
fig. i ; plate 4, fig. 12.

Umbrella about 3.5 mm. in width and height. Somewhat bell-shaped with fairly thick
walls. 4 marginal, long tentacles with very large, spear-head-shaped, basal bulbs one-third as
lung as the bell-height. The ectoderm of the tentacles is thickly crowded with nematocysts.
The entoderm of the tentacles and basal bulbs is pigmented. Velum narrow. 4 wide, straight
radial-canals. Stomach about as wide as long placed upon a short peduncle and extending
a little more than half-way down the cavity of the bell. A row of 50 to fto short, equalh
spaced oral tentacles, each ending in a knob of nematocysts. Gonads form 4 (perradial ?)
swellings extending the entire length of the stomach. Color ( ?) Coast of Ceylon; Chilaw
Paar, and Cheval Paar; March to November.

Genus PODOCORYNE Sars, 1846.

( >)l)\smtirfho;a, PIIILIPPI, 1842, Archiv. fiir Naturgesch., Jalirg. 8, Bd. i. p. ',7, t.if. i, tic,. 3.

I'aitarortnr, SARS, 1846, Fauna littor. Nurveg., tome I, p. 4, t.if. I, figs. 7-18. Hl\< ks, iXdS, British Hvilroul /.,„, phvtes, p.

29.— ALLMAN, 1871, Monograph Tubul. Hydroids, p. 348. Hi MIM., iK'jj. |<>nm. M.nplml., vol. 9, p. 20^. HMtiixir.,

1905, Znolog. Jahrbuchern, Supplement 6, p. 523.

l'u,lm-or\na, KROHN, 1851, Arch, fiir Naturges., Jahrg. 17, Bil. I, p. 263.
Dvimorpliosa, Ai.^si/, A., 1865, North Amrr. A. ,il., p. id}. I'm N. 1X9^, Bihli.itlu-ta X.n.il., Hi-ft n), I.fi;. I, pp. 12, ;;.

MAAS, 1905, Craspcdotcn M.'iluscn ilor fiibo^a KxpcJitmn, Mnnni;. 10, p. 8. V«MIM> » > N, |S.||. /...ol. An/i-ri;iT, Bil. 14,

p. 444. — BEUOT, 1905, Rcvur Suisse dc Zo..]., tmn<- i ;, p. 7-, ; I'-niotorynt, Ibid., p. 102 (literature to 1850). — MAAS, 1906,

Revue Smssr .It- /.mil., form- 14, p. 86, plate }, fit^. 8.

Dysmorphoso+ Cyteeandra, HAKCKKI-, 1879, Syst. iler Medusen, p[i. 76, 79.
Tuiritopsis, AI^ASSIZ, A., 1865, Nnrth Amer. Acal., p. 167.

The name l)\<snitirf>litis<i was first used In Philippi, 1^42, in his description ol I). <on-
<-/i/t olt/, a hydroid polyp which may have been either a Podocoryne or a Hydr actinia in the
modern sense. In 1846, Sars described a closely related, medusa-bearing hydroid under the
name Podocorvne cornea. The name Dysmorphosa was first applied to the free medusa by A.
Agassiz, 1865. The great majority of authors have included both the hydroid and the tree
medusa under the generic name I'oJm <>i \n< . In the absence of proof as to whether Philippi's
hydroid develops medusae or not, we are unable to decide with certainty which name should
stand, but there is no probability that Philippi's hydroid will ever be determined and we niav
avoid confusion by considering it obsolete.

l.'JG MIClirS.K I IF THK WORLD.

GENERIC CHARACTERS.

Margelinae with 4 or more simple, unbranched oral tentacles and with 8 or more simple
marginal tentacles. When present the peduncle of the manubrium is simple, solid, and gelat-
inous. The hydroid stock is Podocoryne.

Haeckel, 1879, restricts the genus "Dysmorphosa" to include only those medusae which
have 8 marginal tentacles. Medusze bearing more than 8 tentacles he places in a new genus
"(1 \iirunJi n." Inasmuch as specimens of Podocoryne fulgurans and P. cnrnea are often found
which have more than 8 tentacles, we consider Haeckel's definitions to be too restrictive. Van-
hoffen, 1891, defines Dysmorphosa, as we do Podocoryne, to include medusa, with 8 or more
marginal tentacles.

PoJ'jiftr \iii- may be distinguished from Lymtiorca by its simple unbranched oral tentacles.
In Turritopsis there are no oral tentacles, but only 4 simple lips studded with nematocyst-
knobs, and the presence of simple oral tentacles in Podocoryne at once distinguishes it from
Turritopsis. It is distinguished from Cyta-is by its having eight or more marginal tentacles,
whereas C ytieis is restricted to medusae bearing 4 marginal tentacles. In Cvtccis the oral ten-
tacles are commonly more than 4, whereas there are only 3 known species of Dysniorp/ios/i
bearing more than 4 oral tentacles. Podocoryne is possibly derived from the more simply
organized C \ta-is.

Podocoryne carnea Sars.

Plate 14, figs. 2 to 6; plate 15, fig. 14.
SYNONYMS OK THK EUROPEAN FORM.

( •) Dysmorphosa conchicola, PHILIPPI, 1X42, Archiv. fiir Naturgesch., Jahrg. 8, Bd. i, p. 37, taf. i, fig. 3.

Podocoryne carnra, SAKS, 1846, Fauna Littor. Norveg., p. 4, taf. i, figs. 7-18.— KROHN, 1851, Archiv. fiir Naturgesch., Jahrg.
17, Bd. I, p. 266. — HINCKS, 1868, British Hydroid Zooph., p. 29, plate 5, 6 figs. — ALLMAN, 1871, Monograph Tubul. Hy-
droids. p. 349, plate 16, figs. 1-9. — DEVARENNE, 1882, Archiv. de Zool. Ejpcr., tome lo, pp. 645, 674, 683, plate 33, figs.
6-15; plates 34, 36-38; 1882, Compt. rend. Paris, tome 94, p. 892. — WF.ISMANN, 1883, Seiualzellen bei den Hydromedu-
sen, pp. 63-72, taf. 19, fign. 1-13. — BROWNK, 1896, Proc. Zool. Soc. London, p. 463. — DEVARENNE, 1881, Annals and
Mag. Nat. Hist., ser. 5, vol. 9, p. 134.— HAMANN, 1882, Jena. Zeitschrift fur Naturwissen., Bd. 15, p. 517, taf. 20, fign. I,
3, 4(histology of hydroid). — JICKELI, 1883, Morphol. Jahrb., Bd. 8. p. 621, taf. 27 (histology of the hydroid). — GRAEFFE,
1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 347. — ISHIKAWA, 1888, Zeit. fiir wissen. Zool., Bd. 47, p. 621, 6 fign. (origin nf
egg cells). — BEDOT, 1905, Revue Suisse de Zoo]., tome 13, p. 103 (literature 1842-1850). — RITCHIE, 1907, Trans. Roy.
Soc. Edinburgh, vol. 45, p. 1^23 (from Cape Colony, South Africa).

Dysmorphosa carnea, HAECKEL, 1879, Syst. der Medusen, p. 77.

(:\[,ti! exi^nti, HAECKEL, Ibid., p. 634.

Podocoryne h'fckcln, HAMANN, 1882, Jena. Zeit. fur Naturwissen., Bd. 15, p. 519 (young stocks of I', carnea ?).

(?) Podocoryne conchicola (Philippi) in part, HARGITT, 1904, Mitth. Zool. Station Neapel, Bd. 16, p. c8l, fig. 26, taf. 22.

SYNONYMS OF THE AMFRICAN REPRESENTATIVE.

Turritopsis nutricula, AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 97, figs. 22, 23; 1865, North Amer. Acal.,p. 167*
figs. 269, 270. — FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 153, plate 4, figs. 4, 7-10.

t'.nli \<tion formosuii, FEWKES, 1882, Bull. Mus. Comp. Zool., vol. 9, p. 294.

Podocoryne carnpa, BUNTING, 1894, Journ. Morphol., vol. 9, p. 205, plate I I, fig. 68. — LhviNSEN, 1893, Vid. Meddel. Nat. For.
Kjhbenhavn (5), Bd. 4, p. t 53.— HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 38, plate 4, fig. 5 (figure
labeled " Turritopsis nutricula," on p. 37).

Turritopsis nutricula (in part), NETTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 375.

Porocoryne carnea, HARGITT, 1901, American Naturalist, vol. 35, p. 582, fig. 44.

Podocoryne, HAZEN, 1902, Amer. Naturalist, vol. 36, p. 193 (regeneration).

AMERICAN VARIETY.

Adult nii-i/iisa (plate 15, fig. 14). — Bell ellipsoidal in form and about 3.5 mm. in height.
Gelatinous substance not very thick, but quite tough and rigid. There are about 24 to 32 mar-
ginal tentacles which are about as long as the bell-height. The tentacles- are not very flexible,
and are usually carried curled upward. The tentacle-bulbs are well developed and are filled
with entodermal pigment granules. The velum is well developed. There are 4 straight
and narrow radial-canals. The manubrium is flask-shaped and there is no peduncle. The
mouth-opening is surrounded by 4 short, radially situated, unbranched, oral tentacles, each of
which terminates in a knob-shaped cluster of nematocysts. The ripe ova and spermatozoa are
found in the 4 interradii within the ectoderm of the manuhrium. According to Ishikawa and

PLATE 14.

Fig. I. PoJocorynr niinuta, Tortugas, Florida, July 16, 1898.

Fig. 2. The hydroid of Podocoryne cornea. Agassiz Laboratory, Newport,
Rhode Island, June 20, 1895.

Figs. 3 to 5. Podocoryne carnea. Stages in the growth of the medusa. Agas-
siz Laboratory, Newport, Rhode Island. July 20 to 24, 1892.

Fig. 6. Podocoryne carnea, young medusa. Agassiz Laboratory Newport,
Rhode Island, July, 1895.

Fig. 7. PoJocor\'iii' duhia. Tortugas, Florida, July 20, 1898.

Fig. 8. Podocoryne diibui. Side view of oral tentacles.

Fig. 9. Podocoryne Jubia. Side view of marginal tentacles.

Fig. 10. Tumtopsis nutricula, mature male. Agassiz Laboratory, Newport,
Rhode Island, July, 1892.

Figs. II to 13. Tumtopsis nutncula, young stages of the medusa. Agassiz
Laboratory, Newport, Rhode Island, July, 1892.

Drawn from life, by the author.

PLATE 14

•

AVI'lluMI-'.DI'S.K roh KYNK.

between European and American meiiusiT of Po<locor\rif r,ir'ifti.

Podocoryne carnea Podocorynr i.irnra
Sars, of Europe. var. americana.

Bunting, the ova originate in the entoderm, Init the sperm originates and remains in the ecto-
derm. When the medusa is mature both ova and sperm arc found in the ectoderm of the
stomach. In the female, the ova are large and prominent and are spherical in form. The
entoderm of the manubrium and of the tentacle-bulbs is red, or brown to red, in color.

H yJroiJ unJ \oung medusa. — No specific difference exists between the American and
European hydroid stocks of Podocoryne, The hydroid (plate 14, fig. 2) is commonly found
upon shells which are tenanted by the hermit-ciah (I'lignnn) and also upon the carapace of
I.imulus. The polypites arise at somewhat irregular intervals from a hydrorhi/.a which clings
to the surface of the shell, etc., upon which the stock is growing. In young colonies the hydio-
rhiza consists of an open network of anastomosing fibers, which are covered by :i thin, delicate
pensarc, and externally by a fleshy hydrocaulus, or ccrnosarc. As the colony becomes older,
however, the fibers of the hydrorhiza form a closer network, and the chitinous perisarc fills in
the spaces between them. Numerous short, chitinous spines are developed upon the crust
which covers the fibers, and thus we find the polypites arising from a fiat, spinous base which
adheres to the surface on which the colony is growing. The colony is composed of two kinds
of hydranths: sterile feeding-polypites, and reproductive gonostyles. The sterile feeding-
polypites are spindle-shaped, being about twice as wide near the oral circlet of tentacles as
they are at the base. They have 12 to 16 straight, stiff" tentacles. The mouth is situated at the
apex of a dome-shaped proboscis. They are very contractile and may vary in length from about
5 to I 5 mm. The reproductive polypites, or gonostyles, are frequently exactly similar in si/.e
and shape to the feeding-polypites, and, in tact, are probably merely feeding-polypites which

have developed medusa-buds
(see plate 14, fig. 2, <;). In
other instances the gonostyles

o

are smaller and more slender,
and possess not more than 4
to 8 tentacles (see g', fig. 2>.
The medusa-buds arise from
a zone which is slisrhtlv below

D

the circlet of oral tentacles.
From 4 to 8 of these buds are
usually to be seen upon each
gonostyle. According to Mar-
tha Bunting, 1894, it appears
that the medusa-bud arises as an outpushing of both entoderm and ectoderm of the gonostyle.
As the bud progresses in its development, we find the ova in the entoderm of the manu-
brium. When a later stage has been reached the}' migrate from the entoderm into the
ectoderm. The spermatozoa, on the other hand, originate in the ectoderm of the manu-
brium, as was shown by Weismann, 1883. When set free the medusa usually has 8 ten-
tacles: 4 radial and 4 interradial. The radial tentacles are usually better developed than the
interradial and in some individuals there are but 4 tentacles at the time of liberation, the inter-
radial ones not yet being developed.

It is remarkable that while in some stocks of Podocoryne the medusae are set free in an
immature state, in others the medusae are mature when liberated, the manubrium being
distended with sperm or ova, which are discharged almost immediately after the medusa is
set free. It is possible, as Allman, [871, suggests, that this difference may be due to the influ-
ence of local conditions, which may be favorable in the one case and not so in the other to an
advanced development of the medusa. Krohn, 1851 , and I. oven, [857, have observed Stocks ol
the European form of PvJticnrvni* which were setting ticc mature medus:e. Indeed, we appeal
to have a parallel case in Stinm on the Massachusetts coast, where immature medusae are liber-
ated during the early spring months, whereas the medusae become ripe, discharge then genital
products and wither upon the hydroid stock in May. Good figures of I'lnlmm \n<- stocks which
are setting free immature medusae have been given by Sars, 1846; Ilincks. ifsViS; Allman, iX-i;
etc. When set free the medusae commonly have 8 tentacles, 4 radial and 4 interradial. The
radial tentacles are usually more advanced than the interradial, and in some few individuals

Number of tentacles when liberated
from hvdroul.

Number of tentacles possessed bv
mature medusa.

Mrilus.i-lnuls on interradi.il suit's i>f
stomach of medusa.

Usually 4 to 8, rarely
ih (Graeffe).

Observed by Sars,
1846.

4 in S. Usually 8.

16 to 32. I'su.illv

.iln.ut 24.

No medusa-buds
observed .

138

MF.ni'S.K OF THK WORLD.

there is no trace of interradial tentacles at the time when the medusa is liberated. The manu-
hrium is short and fusiform and the mouth is surrounded by 4 radially situated, oral tentacles,
each of which terminates in a knob-shaped cluster of nematocysts. When set free the medusa
is about 0.5 to 0.6 mm. in height.

In those medusae which are set free in an immature condition there are at first 8 tentacles,
but these increase in number as growth proceeds and finally, when the medusa is about 3.5 mm.
in height, there are usually about 32 tentacles, 8 in each quadrant. The manubrium of the
young," immature medusa is slender and fusiform, while in those medusae which are set free in a

I'u.. 74.— /

c

- tii'iii-ii, livdroid ami young medusa. Drawn from lift" by Professor William K. Brooks
and kindly presented to the author for publication in this work.

mature state it is globular and greatly distended with the genital products. The ectoderm of
the hydroid is slightly bluish and translucent, while the entoderm is creamy-pink, or silvery-
white in color.

This species has been found upon the Atlantic coasts ot Europe, in the Mediterranean
Sea, and trom Saldanha Bay, Cape Colony, South Africa. Levinsen, 1893, records it from the
west coast of Greenland, and we have found it in great abundance in Narraganseit and
Buzzard's Bays, on the southern coast of New England. It has not been taken at Beaufort,
North Carolina, nor at any station farther south. The medusae are very common in Narra-
gansett Bay from the middle of June until October, and I enjoyed exceptional opportunities

PLATE 15.

Fig. i. Lvmnorfa borealis. Eastport, Maine, September 19, 1898.

Fig. 2. Lymnorea borealis. Side view ot the oral tentacles.

Fig. }. Lvmnorfa borealis. View ot the nematocyst-capsules upon the oral

tentacles.
Fig. 4. Lymnorea alexandn, young medusa. Nassau Harbor, Bahamas,

June, 1903.
Fig. 5. Lymnorea alexandn, mature medusa. Nassau Harbor, Bahamas,

July, 1903.
Fig. 6. Lvmnorea alexandri. Cross-section showing the gland-cells ot" the

mid-region ot a radial-canal of the medusa shown in figure 5.
Fig. ". Lvmnorfa alexandrl. Knob-like end ot" one of the oral tentacles

showing the nematocysts.
Fig. 8. Lvmnorea ale\anJri. Mouth parts ot" the young medusa shown in

figure 4.

Fig. 9. Lvmnorea ale\andrt. Marginal tentacle.

Fig. 10. Turritopsis nutricula, young medusa. Agassiz Laboratory, New-
port, Rhode Island, August, 1892.

Fig. ii. Turritopsis nutricula, young medusa. Newport, Rhode Island.
Fig. 12. Turritopsis nutricula. View ot the lips ot a mature male medusa.

Tortugas, Florida, April 6, 1906.
Fig. 1 3. Turritopsis nutricula. Aboral view ot manubrium showing the

vacuolated entodermal cells surrounding radial-canals. Tortugas,

Florida, April n, 1906.
Fig. 14. Podocoryne carnea, mature female. Agassiz Laboratory, Newport,

Rhode Island, July, 1895.

Drawn trom lite, by the author.

PLATE 15

10

13

AXTHflMKDrS.F. — PODOrORYXE. 130

for observing their growth and development while studying at Dr. Alexander Agassiz's labor-
atory at Newport, in 1892-96.

In 188 1-82 de \ arenne concluded that both eggs and sperm-cells originate in the entoderm
of the crenosarc of the hydroid and afterwards migrate into the medusa-bud. The more
careful researches ot Weismann, 1883, however, refuted this view, showing that the male
germ-cells originate in the ectoderm of the budding medusa, but do not wander from their
place of origin. On the other hand, the female germ-cells may possibly originate in the ecto-
derm; if so they soon wander into the entoderm of the budding gonophore, then into the spadix
of the medusa-bud and finally into the ectoderm of the manubrium of the medusa. According
tii rlie later studies of Ishikawa and of Bunting, however, the egg-cells t,ri^inntf in the min-
der m dl the medusa-bud and afterwards migrate into the ectoderm.

Plate 12, figs. 5 to 9; plate 11, figs, i to 5.

Adult medusa. — Bell usually somewhat pynform in shape and about I mm. in height.
Gelatinous substance very soft and flexible, so much so, indeed, that the medusa is often found
to be much distorted and shriveled by the powerful contractions of the circular muscles. Very
frequently this contraction causes deep interradial furrows to appear upon the surface of the
bell, in the meridian of the radiating muscles of the subumbrella, and I believe that this
appearance led Fewkes, 1881, to describe one ot these contracted medusae as a new genus:
"Mabella gracilis." There are usually 8, but sometimes 16 (plate 13, fig. 5), long, marginal
tentacles with well-developed basal bulbs. These tentacles are quite stift and are carried
curled upward. The tentacle-bulbs are large and are filled with entodermal pigment. The
velum is wide. There are 4 straight, slender, radial tubes. The manubrium is furnished
with a well-developed peduncle (plate 12, figs. 6 to 9), which is quadratic in cross-section.
The peduncle has the shape of a frustum of a pyramid, and it extends about one-third the
distance from the inner apex of the bell-cavity toward the velar opening. The 4 radial tubes
extend down the 4 angles of the pyramid into the gastric part of the manubrium. The gastric-
portion of the manuhrium is short and the mouth-opening is situated at the lower extremity of
a short tubular neck. There are no prominent lips. 4 radially situated, oral tentacles arise
from the sides of the neck. These oral tentacles are quite flexible. They taper gradually and
end each in a single, knob-shaped cluster of nematocyst-cells. Medusa-buds are produced
upon the sides ot the stomach. These buds arise in the interradn (plate 12, tig. 7), and one
usually sees 4 or 5 of them in various stages of development. The place and time of appear-
ance of these medusa-buds appear to follow the law demonstrated by Chun, 1895, in Rnthkta
ni tripnin'tata. When set free the young medusa has 8 marginal tentacles. It is interesting to
observe that while the medusae are still attached to the manubrium of the mother they
themselves begin to develop medusa-buds upon the sides of their manubria (plate 12, tig. 7).
The entoderm of the tentacle-bulbs is of a delicate salmon-red. The manubrium is some-
times colorless and sometimes salmon-red.

This medusa is very abundant at Newport, Rhode Island, late in the summer, when it
occurs in vast swarms which cause the water, when disturbed at night, to glow with ;i bluish
phosphorescence. A large swarm was found in the St. Mary's River, Maryland, near the
mouth of the Potomac, in November, 1905; and I have also taken it at Beaufort, North Caro-
lina, late in November. It has not been seen at Tortugas, Florida,

This species is readily distinguished from Podocoryne rarnea by its much smaller size.
Also the manubrium is provided with a well-developed peduncle, while in I1, ctirnnt there is no
peduncle. P. fiilgnrntis is very delicate and flexible, while P. faniea is tough and relatively
inflexible,

A PoJdiat \iii- found at Beaufort, North Carolina, by Professor Brooks, is probably the
hydroid of this medusa. The tentacles are usually in whorls of 5.

1 |() MEDUS.E OF THE WORLD.

Podocoryne areolata Hincks.

Hvilractinia areolata, ALDER, 1861, Trans. TynesiJe Nat. Field Club., vol. 5, p. 225, plate 9, figs. 1-4 (hydroid).
Podocorme areolata, HINCKS, 1868, Hist. British Hydroid Zooph., p. 32, plate 6, fig. I.
C.\'t.eanilra areolata, HAECKEL, 1879, Syst. der Medusen, p. 79.

I I C.\titandra areolata, BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, 'p. 262; 1897, Proc. Zool. Soc. London, p. 817,
plate 48, figs. 1-2; 1903, Bergens Museums Aarbog, No. 4, p. 12.

This medusa is liberated from Podocoryne areolata of the British coasts.

According to Browne, the medusa becomes about 4 mm. wide and slightly less in height.
The bell is pyriform with a thick, gelatinous apex. There are 25 to 30 marginal tentacles
about as long as the bell-radius and with fairly large basal bulbs which lack ectodermal
ocelli, but have entodermal pigment. The stomach lacks a peduncle and is about one-halt
to two-thirds as long as the depth of the bell-cavity. There are 4 long, narrow lips, each of
which bifurcates and ends in 2 knob-like clusters of nematocysts. The gonads are upon the
sides of the stomach.

According to Browne, the entoderm of the stomach and tentacle-bulbs is rich reddish-
brown, darker than the stomach.

When set free from the hydrnid the medusa has 16 marginal tentacles of various lengths.
Browne has not observed the hydroid, which was obtained by Alder. I am inclined to believe
that this medusa may prove to be identical with Lymnorea borealis, found at Eastport, Maine,
on the American Atlantic coast. In Europe it ranges from British coasts to Norway, and is rare
according to Browne.

Podocoryne octostyla.
D\smorpliosa oe/ostvla, HAECKEL, 1879, Syst. der Medusen, p. 78, taf. 6, fig. 6.

Bell 0.4 mm. high, 0.5 mm. wide, with bulging sides and low, conical apex. 8 short,
equally developed, marginal tentacles, about as long as the radius of the bell, with small basal
bulbs. Stomach mounted upon a conical peduncle as long as itself; both together about
two-thirds as long as the depth of the bell-cavity. 8 simple, knobbed, oral tentacles about as
long as the stomach. 4 swollen, interradial gonads on the sides of the stomach. When young,
medusa-buds are produced upon the interradial sides of the stomach. Gonads and tentacle-
bulbs nearly black.

Found in the Mediterranean. Two specimens were found by Haeckel.

Podocoryne polystyla.
Cytaandra pol\st\la, HAECKEL, 1879, Syst. der Medusen, p. 79.

Bell nearly cubical, 1.5 mm. wide, 1.5 mm. high. 32 short, marginal tentacles (without
ectodermal ocelli ?). Stomach large, cubical, mounted on a short peduncle and filling nearly
all of the bell-cavity. 16 short, simple, oral tentacles. 4 gonads on the lower part of the
stomach-wall. The gonads are " halt-moon-shaped " with the convex side below. Stomach
yellow. Gonads and tentacle-bulbs reddish-brown.

Found by Haeckel at Croisic, Atlantic coast of France.

Podocoryne minuta.
Plate 14, fig. i.

nismorpltosa minuta, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 41, plate 18, fig. 42.

Dvstnorpltosa minuta, TRINCI, 1904, Monit. Zool. Ital., Anno 15, p. 304.

( ) ( ,r.F/s minima, TRINCI, 1903, Mittheil. Zool. Sta. Neapel, Bd. 16, p. I, taf. I, fign. 1-30 (young medusa ?).

Bell pear-shaped and only 0.3 mm. high. Bell-walls quite thick and the apex solid and
bluntly pointed. In common with other species of Podocoryne, the gelatinous substance
of the bell is very delicate and the medusa soon collapses into a shapeless mass when subjected
to the abnormal conditions of captivity in a small aquarium. There are 8 solid, marginal
tentacles with well-developed basal bulbs without ocelli. The velum is narrow. There
are 4 straight, narrow radial-canals and a slender ring-canal. The manuhrium is mounted
upon a short, wide peduncle, which is solid, and down the 4 radial corners of which the radial-
canals extend into the stomach. The gastric portion of the manubrium is 4-sided in cross-
section. 4 well-developed, oral tentacles surround the mouth. The entodermal cores of
these oral tentacles are composed of chordate cells, and each tentacle terminates in a bristling.

PLATE 16.

Figs, i to 3. Podocoryne gracilis. Successive stages in the growth of the

medusa. Tortugas, Florida, June, 1899.
Figs. 4 to 5. Bougainvillia autitninalis, young and mature stages ot the

medusa. Agassiz Laboratory, Newport, Rhode Island, Septem-
ber, 1895 and July, 1896.
Fig. 6. Bougainvillia fronJosa, showing planula; upon the sides of the

stomach. Tortugas, Florida, June, 1897.
Fig. 7. Koii^nlnvilliti carolinensis, young medusa. Agassiz Laboratory,

Newport, Rhode Island, July, 1892.
Fig. 8. Bougainvillia carolinensis. Cluster ot marginal tentacles of a

young medusa from Newport, Rhode Island.
Fig. 9. Bougainvillia carolinensis. Oral tentacle of a young medusa from

Newport, Rhode Island.

Drawn from life, by the author.

PLATE 16

AXTHO.MKIH S.t — PODOCOK YNK. 141

knob-like cluster ot nematocysts. Several medusa-buds in various stages of development
are found upon the sides of the upper interradial regions of the stomach. These budded
medusae are set free as in P. fulgurans.

In some specimens the entoderm of the manubrium and tentacle-bulbs is turquoise-blue
and in others it is lilac.

This medusa was abundant at Tortugas, Florida, in the middle of July, 1898, and Trinci
finds a very similar medusa at Naples, Italy. I am inclined to regard Trinci's C.\t,i-n minima
as the immature, 4-tentacled, medusa-budding stage of Podocoryne niniuta. According to
Trinci the bell is pyriform, 0.27 to 0.33 mm. high and 0.2 to 0.24 mm. wide. 4 perradial,
marginal tentacles, each as long as the hell-diameter, with well-developed basal bulbs without
ectodermal ocelli. Manubrium and tentacles yellowish-brown with blue in the entoderm.
Medusa-buds develop upon the 4 interradial sides of the manubrium. These budding medusae
are formed entirely from the ectoderm of the parent medusa.

Trinci in the following year found a medusa similar to the above, but with 8 marginal
tentacles.. Excepting for a slight color difference this 8-tentacled medusa is identical with
I', mi nut a from Tortugas, Florida.

Podocoryne tenuis.

Dysmorphosa tenuis, BROWNK, 1901, Annah and Mag. Nat. Hist., ser. 7, vol. 9, p. 277.

(? ?) Podocor ynt liumilis (hydroid), HARTLAVR, 1905, Zoning. Jahrbuchern, Suppl. 6, p. 523, fig. i.

Umbrella somewhat conical, 2 mm. high and 1.5 wide. There is a slight constriction
above the level of the subumbrella cavity. 8 tentacles. Manubrium cubical and mounted
upon a peduncle about as long as itself. Mouth with 4 lips having terminal clusters of nemato-
cysts. Medusa-buds are found upon the interradial sides of the manubrium. (ionads not
developed in the two specimens studied by Browne. Color ( ?)

Found at Stanley Harbor, Falkland Islands. Briefly described without figures by Browne.
This medusa appears to differ widely from the North American variety of Podocoryne
i ni urn, but the hydroid described by Hartlaub bears a close resemblance to our North Ameri-
can P. canifii. Hartlaub's hydroid may not be the stock of P. trntit>. however.

Podocoryne dubia Mayer.

Plate 14, figs. 7 to 9.
D \smor phasa Jubiu, MAYLR, 1900, Bull. Mus. Cuinp. Zool. at Harvard College, vol. 37, p. 40, plate 21, figs. (14-66.

Bell higher than a hemisphere, 1.5 mm. high and three-fourths-egg-shaped, with thin,
uniform, flexible walls. There are 8 stiff", marginal tentacles which are club-shaped, being
somewhat wider near their ends than they are near their basal bulbs. The outer extremities
of the tentacles are thickly covered with nematocysts. Their basal bulbs are large and swollen,
and a very large, black ocellus is found in the ectoderm ot the under side of each bulb (plate 14,
fig. 9). The velum is well-developed. There are 4 straight, narrow radial-canals and a ring-
canal. At the middle point in the length of each radial-canal there is a small, hernia-like
swelling, which is probably glandular and similar in structure to the swollen parts of the
radial-canals of Lymnorea alexanJn.

The manubrium is pear-shaped, quite slender, and extends about half the distance from
the inner apex of the bell-cavity to the velar opening. The mouth is surrounded by 4 short,
radially situated, oral tentacles, each of which terminates in a bristling cluster ot nematocysts
(plate 14, fig. 8). Cionads ( ?) The entoderm ot the manubrium and tentacle-bulbs is
delicate custard-yellow. Only one specimen has been found. This was taken on the surface
at Tortugas, Florida, on July 20, 1898.

Podocoryne gracilis.
Plate 1 6, figs, i to 3.

C\i.rii gracilis, IWAYtR, njoo, Bull. Mu^. I'oinp. Zool. at Harvard College-, vol. 37, p. -,'i, plat.- -,'.. tig' . 11: 1:4.
Dysmwphosa ^rarilis, MAAS, 1905, Craspedoten Medusen der 5jfro£d Expedition, Monog. 10, p. 8.

Matur,- iiifJusa. — Bell pyriform or dome-like with a slight apical pro|ection. It is about
$ mm. in diameter and a little broader than it is high. The gelatinous substance is of only
moderate thickness on the sides, although it is about twice this thickness at the apex of the

142 MEDUS.E OF THE WOULD.

bell. There are 8 stiff marginal tentacles with large basal bulbs. These tentacles are carried
curled upward from the bell-margin. The 4 radially situated, marginal tentacles are about
two-thirds as long as the bell-height, but the 4 interradial ones are only about halt of this
length. The basal bulbs of all of the tentacles are large, swollen, and hollow, with their
entoderm deeply pigmented. The velum is broad. There are 4 simple, straight radial-canals
and a simple circular vessel. The stomach is mounted upon a short, wide, solid peduncle.
The stomach and mouth-parts are together only about half as long as the depth of the bell-
cavity. The mouth is a simple, round opening without prominent lips and is surrounded by
8 unbranched, oral tentacles, each of which terminates in a knob-like bristle of nematocysts.
4 of these oral tentacles are radial and 4 interradial in position. Their entodermal cells are
chordate, and the nematocyst-capsules upon their free ends are spindle-shaped. Medusa-
buds develop upon the 4 interradial sides of the stomach. When set free each medusa has
only 4 short, equally developed, marginal tentacles. The 4 interradial, marginal tentacles
do not develop until the medusa is about half-grown and after it has acquired 8 oral tentacles.
The entoderm of the manubrium is usually red in color, although often the inner core is red or
pink and the outer annulus of entoderm is green. Each tentacle-bulb is provided with a
dense mass of red, entodermal pigment, which in the case of the radial tentacle-bulbs extends
a considerable distance up the radial-canals. This medusa was quite common on the surface
at Tortugas, Florida, in June, 1899 and 1907.

This species and P. octostyla Haeckel are the only forms of Podocoryne having rrfore
than 4 oral tentacles. The remarkably late development of the 4 interradial, marginal ten-
tacles and the presence of more than 4 oral tentacles indicate the close relationship between
C,\ticis and Podocoryne. In Cyttns more than 4 oral tentacles are the rule. Although the
medusa appears to begin life with but 4, later it acquires 8 or more, Cyta-is vulgans having 32
tentacles.

Podocoryne humilis Hartlauh.

Podocor\ne liutnilis, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 523, fig. K.

( ?) D\stnorphosa tenuis (medusa), BROWNE, 1902, Annal. and Mag. Nat. Hist., ser. 7, vol. 9, p. 277.

Hartlaub describes this hydioid from Terra del Fuego, and states that it may prove
to be the stock of Podocoryne tenuis, which was briefly described by Browne, 1902, from the
Falkland Islands.

The polypites arise in groups from the hydrorhiza. The feeding polypites are about i
mm. long, club-shaped, with narrow bases, and a short, conical hypostome, which is encircled
by a single zone of about 12 filiform tentacles about half as long as the body of the polypite.

The reproductive polypites are swollen, with only about 5 well-developed tentacles
with a cylindrical proboscis, and with 3 to 4 large medusa-buds upon the basal half of the
polypite below the zone of tentacles. These medusa-buds are mounted upon short pedicles,
and when set free the medusae have no sexual products. The hydrorhiza forms an open net-
work of thin branches bearing short, smooth, thorn-like processes. There are no spiral
zooids or tentacular filaments. Color ( ?) Number of tentacles possessed by the medusa
when set free ?

Puerto Bridges, southern Terra del Fuego, January, 1893; from a depth of 7 fathoms.

This species appears to me to be closely allied to, if not identical with, the Podocoryne
of the coast of New England. The number of the tentacles is very variable in our New England
Podocoryne, especially in the reproductive polypites, and the colonies range considerably in
size, apparently being influenced in this respect by conditions of nutrition. There is nothing
in Hartlaub's description to prevent our considering the species from the southern end of
South America to be identical with P. cornea of the coast of New England.

Podocoryne denhami Thornely.
Podocoryne denhami, THORNELV, 1905, Roy. Soc. Report Ceylon Pearl Oyster Fisheries, vol. 2, p. no, plate I, fig. 6.

Basal crust beset with numerous tall, stout, linear, reddish spines. Hydranths white,
with about 24 tentacles on the barren ones, and only 4 or 5 on those bearing gonophores; and
these latter are considerably swollen. A pair of large globule (medusiform ?) gonophores on
each reproductive hydranth. Medusa unknown. Size ( ?)

Growing on a shell of Murc\ in Polk Strait, Ceylon.

ANTIIOMKDI S.K— -TVHKITol'SIS. I I.'!

Genus TURRITOPSIS McCrady, 1856.

OfCiiniti (Turritopsisjj M<C'RADY, 1856, Proc. Elliott Soc. Charleston, p. i.

MtiJcrriii, FKWKKS, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 149.

Turrilofiis, MI-CRA.DY, 1857, Gymn. Charleston Harbor, p. 24.— HAI.C-KM., 18^9, Svst. d.-r Medusen, p. 65.— BROOKS. i*S;.
.Studies Biol. Lab. Johns Hopkins Univ., vol. 2, p. 465. — BROOKS, 1886, Mem. Boston Soc. Nat. Hist., vol. 3, p. 387. —
HARTLAIJB, 1897, Meeresuntcrsuch., Helgoland, Neue Folge, Bd. 2, Heft, i, Abt. 2, p. 480.— VAMHHM •>, iXiji. /,.,,il.
Anzeiger, Bd. 14, p. 444.— BROOKS and RITTENHOI si, 1907, Proc. Boston Soc. Nat. Hist., vol. ;-,, p. 4;!..

\on Turritopsis, AGASSIZ, A., 1865, North Amer. Acal.. p. 167.

The type species of this genus is T . nutrient a McCrady, 1856 -57, from Charleston I larlior.
South Carolina. It is common in summer from the West Indies to the southern coast of New
England.

GENERIC CHARACTERS.

Margelinae with S or more simple, marginal tentacles, which arise singly, not in clusters,
from the bell-margin. The walls ot the 4 radial-canals, above the stomach, consist of highly-
vacuolated, entodermal cells, forming a peduncle for the gastric part of the manubrium. The 4
lips are studded with nematocyst-hearing knobs. The gonads are developed upon the 4 inter-
radial sides of the stomach. The hydroid is Dendroclava.

Turritopsis is peculiar in the remarkable development of vacuolated, entodermal cells
lining the courses of the 4 radial-canals above the stomach. This condition is also displayed
to a limited degree by some, but not by all, ot the medusae of the genus Lymnorea. In Lv»i-
nnrt'ii, however, the mouth is surrounded by branched, oral tentacles, whereas in Titrritnf»i>
the lips display bead-like knobs of nematocysts, which I am inclined to regard as the remnants
of the oral tentacles of the ancestral forms of Turritopsis. The tentacles of Turritopsis arc
solid and similar in structure to those ot Lymnorea. There is an ectodermal ocellus upon the
inner (axial) side ot each tentacle, adjacent to the basal bulb. In Tiannje, on the other hand,
when ocelli are present they are found upon the outer fabaxial ) sides ot the tentacle-bulbs. On
account of the close similarity between the tentacles of Turritopsis and those of (),,,inin,
Lymnorea, Podocoryne, Cytteis, znd Cubogaster, we incline to Vanhbffen's (1891) view that
T urritopsis is related to the Margelinae rather than to the Tiarina.-.

Turritopsis is closely related to Oceania, but is distinguished by its \acuolated peduncle,
that of Ofcanta being solid and gelatinous.

Turritopsis nutricula McCrady.

Plates 14 and 15, figs. 10 to I}.

Oftania (Turritopsis) tiutricula, McCfiAuv, 1856, Proc. Elliott Soc. Charleston, pp. i-;6, plate 4, figs, i -10.

Turritopsis nulricula, MiCRADY, 1857, Gymn. Charleston Harbor, p. 25, plate 8, fig. i .— BROOKS, iSS;, studies Johns Hopl.m
Univ. Biol. Lab., vol. 2, p. 465. — BROOKS 1886, Mem. Boston Soc. Nat. Hist., vol. ; p. ;SS. plate ;?. -HAROITT, 1904,
Bull. I'. S. Bureau of Fisheries, vnl. 24, p. }7 (the figure is I3odo(or\nt earned). — Rm'tNiMH s>, M>O^, Pnn . Ii.i'f.,n s.,, .
Nat. Hist., vol. 33, p. 437, plates 30-35, figs. 1-55 (development).

.U"./<vn<; mullitentacula, FKWKKS, 1881, Bull. Mus. Comp./ool. at Harvard Collrp-, \nl. S, p. 141), pi. tic ^, figs. 7-9.

Mvtlrctiii nutrictiiti, FKWKLS, 1882, Bull. Mus. Comp. 7<ool., \D|. »), p. 2<K-

non Turritofsis nulricula, AI.ASSIZ, A., 1865, North Amer. Acal., p. 167, figs. 269, 270.

Turritopsis nulricula (in part), HAF.fKti., 1879, Syst. der Medusen, p. 66. — NtTTiN(., 1901 , Bull. I . S. Fish C'nnmiss!<.n. \i»l. i •).

P- 375-

EL Roi-h AN MMH SA.

Oti-iinia polvfirr hti, KH-KRSTUN, lSfi2. X,eit. fur \vissen. Xool,, Bd. 12, p. i'i. taf. 2, figii. II i;.

Turritopsis potvnttmi, HAF.CK>L, 187"). S\st. dcr Mi'ilusen. p. (>tt.

Turrilrjptrs f>rjl\'t irrlia, HARTLAUB, 1897, Wis.sen. MecresunterMiili . K "1111111 .. MI rr< Kir I, H<-|L;"|,UI.|. \' ur I "lee, Bd . 2, p. 480,

taf. l6r, fig. 2.
( ) r\/,r/.t /)d/v5/\//;, \\'N.I., 1844, Hone Tergcst in.r, p. 68, taf. 2, fig. V.

.•hncriian IHK/HSH ( plate 14, fig. 10). — Bell usually slightlv pvritorm and about 4 to , mm. in
height. Bell-walls uniformly thin. There are 40 to 70 simple, marginal tentacles, which are all
ot about the same length and are somewhat shorter than the bell-height. T hese tentacles are
capable of much contraction or extension. Their basal bulbs are large and there is a single,
ectodermal pigment-spot upon the lower (centripetal) side of each tentacle near its place of
origin from the basal bulb. The surface of the tentacles i> co\i-ied thickly with nematoc\st-
cells. The velum is well developed. There are 4 straight, narrow radial-canals and a narrow
circular vessel. The manubrium is large and fills about halt of the bell-cavity. The upper

144

MEDUS/E OF THE WORLD.

part of the manubrium near the base consists of 4 radially situated masses of large, highly
vacuolated, entodermal cells, through the midst of which the 4 radial-canals extend downward
into the stomach. These cells are indeed only the entodermal walls of the radial-canals (plate
14, fig. 13). The stomach is large and quadratic in cross-section. The cruciform mouth is
situated at the extremity of a short neck and is surrounded by a row of nematocyst-bearing
knobs (plate 15, fig. 12). The gonads are developed upon the sides ot the stomach, where
they occur in the form of a double, longitudinal, swollen region in each adradius. Their
outer surfaces are smooth. The entoderm of the stomach is dull-yellow or orange, or dull-
vellow streaked with orange. The tentacle-bulbs sometimes contain a little entodermal
orange pigment. The ocelli upon the tentacles are dark-brown or orange.

Hydroid and young medusa. — The hydroid (fig. 76) was found by Brooks, 1883, on piles
of a wharf at Morehead City, North Carolina. It is a Dendroclava and closely related to D.
dohrnii Weismann. The stems of the hydroid are from 8 to 10 mm. in height and bear
large, terminal hydranths. There are also numbers of short, side branches which terminate in
hydranths. The main stem and the side branches are incased in a loose, cylindrical pen-
sarc, which is thick and becomes mcrusted with foreign matter. The pensarc is not an-
nulated, and terminates abruptly by a sharp collar immediately below each hydranth. The
hydranth or feeding-polypite is long and fusiform and bears from 18 to 20 short, thick,
filiform tentacles, which are arranged in three or more indefinite rows or whorls. The medusa-
buds originate upon the sides of the stem at the bases of the hydranths. Each medusa-bud is

borne upon a short stalk or peduncle and is closely
invested by a thin capsule of perisarc. When set free
the young medusa has 8 tentacles. The manubrium is
cone-shaped and there is a large peduncle formed of
highly-vacuolated cells. 4 prominent, nematocyst bear-
ing knobs surround the mouth. The hydranths are pale
yellowish-red.

This medusa is found from the coast ot Cuba to the
southern coast of New England, being about equally
abundant in the northern and southern limits of its range.
It is very common in the Bahamas and at Tortugas,
Florida. In Charleston Harbor, South Carolina, it is
commonly infested by larvae of Cunoctantha octonaria
McCrady, although it appears to be quite immune from
this parasite in other places. I can detect no specific
distinctions between this medusa and "Turntopsis [>ol\-
cirrka," which is occasionally seen off the Atlantic coast
of France and German}'. This form is well figured by
Keferstein, 1862, and Hartlaub, 1897, who are the only European students who have observed
the medusa on the eastern side of the Atlantic.

Rittenhouse, 1907, has made an elaborate study of the development of T. nutricula. The
ova develop in the ectoderm of the 4 interradial sides ot the manubrium. The primitive ova
grow by the absorption ot ovarian cells around them, as is common in other hydromedusae.
The yolk-spheres in the ovum are formed from the ovarian cells which it absorbs. About 20 to
35, rarely 50 or more, eggs are discharged into the water by the muscular rupture of the ovar-
ian walls between 5 and 6 o'clock in the morning. The discharged eggs are spherical, o.i 16
mm. in diameter and have no membrane. They are yellowish-white, heavier than sea-water,
and opaque. The outer layer of finely-granular ectoplasm is distinct from the coarsely gran-
ular, yolk-laden endoplasm. Soon after being discharged the egg gives off two polar bodies
and is fertilized. Segmentation is total and approximately equal. The first two segmentation
planes are meridional and the third equatorial. The blastomeres remain quite far apart,
touching only slightly. After this the cleavage becomes remarkably irregular, recalling the
extraordinary condition observed by Hargitt in Pennaria. A solid morula is formed, which has
no central segmentation cavity and which resembles a loosely-connected mass of irregularly
grouped cells rather than an embryo of any metazoan. The cleavages follow one after another
at intervals of 20 to 30 minutes.

I''K,. 75. -Turrtlopm nutrtculti, after Brook
Mem. Boston Soc. Natural History.

ANTHOMEDUSiE— TUBRITOPSI8.

145

As in MetschnikofF's <>,,,iin<i /inntitn and Margin's embryos ol ' Pi-iiiinrin, this ii regularly
shaped morula gradually changes into an oval embryo, the surface of which becomes ciliated. BO
that it swims upward from the bottom. This takes place at about 4 in the afternoon in eni^
which were laid between 5 and 6 in the morning.

Rittenhouse finds that during this period, when the loose, irregularly shaped mass of cells
shapes itself into an oval embryo, the ectoderm and entoderm are formed. During this period
"the cell-boundaries are lost tor a short time and a syncvtium is formed. This syncytial
structure is crowded with yolk-granules and nuclei are scattered throughout the protoplasm.
The nuclei soon become more numerous near the periphery and the cell-walls begin to appear"

I'll.. 7'i. - Tun l,Vy/iM< tint! i< Illtl, ,lttrr Hri'oks, in Mi'tll. linstnn Sm . N.ll. Hrl. HvJroul .llhi \nnnr ii"i!v^.l.

between the peripheral nuclei. These peripheral cells are to become the ectoderm, which is
soon separated from the inner, structureless mass by the development of the meaoglcea. I his
inner mass afterwards acquires cell-boundaries between its nuclei, and still later a central
cavity, the ccelenteric space, develops; and thus the entoderm is formed. This ccelenteric
cavity does not develop, however, until the larva is 48 to 60 hours old. The s\ IH\ tmm con-
dition in Turritopsil is much more complete than is seen according to Hargitt in Pennaria, 01
in Bougainvillia, according to Gerd, 1892.

llli

MKIH'.S.-E OF THE WORLD.

Fiu. 77.— Turritopsis, f/>. (') = (0ceania polycirrha),
after Keferstein, 1862, Zeit. fur wissen.
Zool., Bd. 12, uf. 12, fig. n.

Rittenhouse finds that during the early stages ot cell-division in Turritopsis the multipli-
cation is solely by mitosis, but that later, when the embryo becomes a mere irregularly arranged
mass of loosely-compacted cells, some of the nuclei divide amitotically.

When about 50 hours old the elongate, oval larva ceases to swim through the water and
settles down upon its side on the bottom. The larva then becomes the hydrorhiza, or root, of

the hydroid, and the first hydranth arises as a bud
from about the middle of its length. The tentacles
of the hydranth develop in indefinite whorls, with
4 tentacles in each whorl, the oldest tentacle being
nearest to the mouth. In the mature hydroid the
tentacles appear to be irregularly scattered rather
than being arranged in whorls.

Turrrtvpsts should he reared under more natural
conditions than those of the ordinary laboratory
in order to determine whether the remarkable, irreg-
ularly formed embryos described by Rittenhouse
be normal or merely the result of pathological
states induced by adverse conditions; but Miss
Beckwith has recently shown that the cleavage of
Pennarta is normally irregular as is described by
Hargitt.

Rittenhouse finds that when the embryo is in the loose-celled, morula stage it may be
divided into two masses, each one of which produces a normal planula larva of small size.

Turritopsis (?j lata von Lendenfeld.

Tuiritafsis lata, VON LENDENFELD, 1884, Proc. Linncan Soc. New South Wales, vol. 9, p. 588, plate 22, figs. 36, 360.
( ? ?) Mrlirerla pleurostoma, PERON ET LESUEUR, 1809, Ann. du Mus. Hist. Nat., Paris, tome 14, p. 353.

Mature medusa. — Bell 3.5 mm. high, 3 mm. wide, with a low, dome-like, apical projection.
60 to 130 marginal tentacles about as long as the bell-height. These have well-developed
basal bulbs, and there is an ocellus on the inner (velar) side of the base of each tentacle. Velum
well developed. 4 broad, straight radial-canals. The manubrium is mounted upon a peduncle,
but von Lendenfeld does not state whether it be composed of vacuolated, entodermal cells or
of the gelatinous substance of the bell, and his figures are too vague to throw any light
upon this point. Until this is settled we can not determine the systematic position of this
medusa. The stomach is spindle-shaped, about two-thirds as long as bell-height. The
mouth does not extend to the velar opening, and there are 4 lancet-shaped lips which are
recurved and one-third as long as the manubrium. These lips have stalked nettling warts
on their margins. There are 4 protruding gonads on the stomach-wall, each one being
one-fourth spherical in shape. Gonads and entoderm of tentacle-bulbs intensely brown.
Ocelli red.

This is the most abundant medusa in Port Jackson, New South Wales, during the Aus-
tralian summer and autumn.

The young medusae have but 4 marginal tentacles.

Genus OCEANIA, sens. Kolliker, Gegenbaur.

Oceania (in part), PERON ET LESUEUR, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, pp. 344-347.

Oceania, KOLLIKER, 1853, Zeit. fiir wissen. Zool., Bd. 4, p. 323. — GEGENBAUR, 18^6, Zeit. fur wissen. Zool., Bd. 8, p. 223. —

KEFERSTEIN UND EHLERS, 1861, Zool. Beitr. Messina, p. 83. — METSCHNIKOFF, 1886, Embryo). Studien an Medusen,

Wlen, pp. 23, 25, etc., to 78.

Turritopsis (in part), HAECKEL, 1879, Syst. der Medusen, p. 65.
(>)Modecria (young medusa), FORBES, 1848, British Naked-eyed Medusa-, p. 70.
Modteria-'r Callitiara, HAECKEL, 1879, Svst. der Medusen, pp. 47, 48.
Occania=CalIiliara, VANHOFFEN, 1892, Verhandl. Gesell. Deutsch. Naturf. Arztc, 64 Vers., Abth. Syst.. p. 121.

The generic name "Oceania" of Peron and Lesueur has no definite modern meaning,
for it was applied to many medusae belonging to the Eucopidae, Tiarinae, Thaumantidae, etc.
In 1862 "Oceania" was restricted by L. Agassiz to apply to a genus of the Eucopidae, but
this same genus had already been named Phialidium by Leuckart, 1856, and thus Agassiz's

ANTHUMKIH > I. <>< KAMA.

genus Oceania loses precedence. The type species of Oceania in the modern sense is O.
ariniita of the Mediterranean. It is possible that one of Peron and Lesueur's descriptions
refers to this medusa (i.e., Oceania flaviJula?), but I am in much doubt upon this point.
We must fall back upon Kolliker, 1855, and Gegenbaur, 1856, for the definition of Oii-nnin
in the modern sense.

Haeckel suppresses (),,-, nun and merges it with T nrritnpsis, but in Oceania the peduncle
is simple, solid, and gelatinous, whereas in McCrady's T nrrit't[»i<; it is composed of large.
vacuolated cells.

GKNI-RK t H \i< UTKRS.

Margelinje with 8 or more marginal tentacles which arise singly from the hell-niaruin.
and are not grouped in isolated clusters. There are ectodermal ocelli on the inner i \elai i
sides of the tentacle-bulbs. The mantibrium is mounted upon a simple, solid, gelatinous
peduncle. The 4 lips lack oral tentacles, but their margins are lined by a row of wait-like
nematocysts. The hydroid is clava-like.

Following the lead ot Agassrz, it has been the custom of American students to define
Oi i uniii in a sense equivalent to PhniliJium of Leuckart, but as we ha\c seen I'/miliJiuin,
1856, clearly takes precedence over Agassiz's Oceania, 1862.

In the system here proposed the genus Oceania falls among the Oceanidx instead of
being among the Eucopidae as it would be were Agassiz's definition admissible.

FIG. 78. — "MoJrrria joTtno'a" after Forbes.

FIG. 79. — Oceania "faviitult" (O . armata), after Gegenhaur, in Zeit. fur «i en. /.<">

nenc name Modeena Forbes, 1848, was applied to a very voi

. y.ooi., Bd. 8.

I he generic name Modecria Forbes, 1848, was applied to a very young and immature
medusa which is so vaguely described that we can not state whether or not it had ectodermal
ocelli. I consider that the name Modecria had best be dropped, having already led to nearly
endless confusion.

The genus Oceania is closely related to Turritopsis, but is distinguished by its simple,
solid, gelatinous peduncle; whereas the peduncle in Turritopsis is composed of a mass of
vacuolated entodermal cells.

Oceania armata Kolliker.

( ?) Moiieertu forrnottl (young mrihisa), l-'oKli^, |S4S. Unti^i Naked-eyed M'- lir,,i', p. 70, pl.itr 7, tii;. I.

Oceania armala, KCILI.IKF.R, 185^, Zoit. fur \visM-n. 7-o.il., BJ. 4. p. -,2',. — M ITSI nMKot \ , iSSd, Kinlirynl. Stiulii-n an Mnlu rn.

\\'ifn, p|i. 2^, 25 (egg); 36, 37, 47 (segmentation); 75, -S (development of hydroid).
Offtinia ftartttula, GEGENBAUR| 1856, Zeit. fur uissen. Xoo] ., It, I. X. p. 12:;, taf. 7, fig. 4. KSJIRMMN i MI l^nifks. 1861,

Zoolog. Beitragc, p. 83, Lrip/ii;. -SpAciNoi.INI, 1876, Atti Soc. Ital. Sci. Nat., tome 19, p. 22, lav. ;, fig. 3.
Pandea fliiridttla, A(;ASSIZ, L., 1862, Cont. Nat. Hi.st. I1. S., vol. 4, p. ^4-. •

Turritopsn armala, HAM'K'F.L, 1879, Syst. der \fetlusen, p. 65.
t'.allitiata polyopthalma, HAM KM, 1X71), Svst. Jer Metlust-n, p. i\-j, taf. 3, fign. 1-5.
Tiarella parthcnnpia, TRINCI, 1906, Monitore Zoologico Ital.. Anno i^, p. ioK, 2 figs.

Bell pyriform, flat-topped, higher than a hemisphere, with uniform, thin walls. 8 to 10
mm. high, 8 to 10 mm. wide, no to 100 or more slender, tapering, marginal tentacles. 1.5
times as long as the bell-diameter. 1 hese tentacles are crowded into a double row. Their

148

MKDUS.E OF THE WORLD.

entoderm is solid and composed of chordate cells. There is an ectodermal ocellus on the velar
side of each tentacle-bulb. Velum well developed. Well-developed circular muscles in the
subumbrella. There are 4 jagged-edged, slender radial-canals and a simple ring-canal.
The stomach is flask-shaped, cruciform in cross-section and mounted upon a short, pyramidal,
4-sided peduncle. This peduncle is a solid, gelatinous, truncated pyramid, only about one-
third to one-fifth as long as the stomach itself. It is not vacuolated as in Turntopsis. The
4 lips are large and recurved and extend to the middle of the bell-cavity and their edges are
studded with a single row of knob-like nematocyst-warts. The 4 gonads are developed upon
the interradial sides of the stomach and are longitudinal swellings with smooth outer surfaces.
The stomach and gonads are yellow or brownish-yellow. Eggs pearly-white. Lips red-
dish-yellow, or port-wine colored. Radial-canals, ring-canal, and tentacles milky-yellow.
Ocelli reddish-brown (see fig. 91).

Found in the Mediterranean. Ouite common at Naples. (See figs. 78-81.)
Vanhoffen (1902, Verhandl. Gesell. Deutsch. Naturf. Arzte, 64 Vers., Abth. Sitz., p. 121)
maintains that this medusa is identical with CuUtttara polyopthalma Haeckel of the Canary
Islands, and with this I am in accord, having seen many specimens of the living medusa at
Naples. Trinci, 1906, describes a medusa from Naples which resembles O. armata in all
respects excepting that he finds no ocelli. He studied preserved specimens, however, and

FIG. 80. — Oceania. "Callitiara polyophlhthalma11 (Oceania armata), after Haeckel, 1879.

the ocelli may have faded. As Gegenbaur states, the ocelli upon the inner sides of the tentacle-
bulbs recall the conditions seen in Bougainvillia, and oblige us to place the medusa among
the Margelinae. In the Tiarinae, on the other hand, the ocelli when present are always on
the outer, abaxial sides of the tentacle-bulbs.

Gegenbaur (1854, Zur Lehre von Generationswechsel, p. 28, taf. 2, fign. 10-16) reared
the larva from the egg, and later Metschnikoft carried out a similar series of studies.

Metschmkoft, 1886, finds that the egg of this medusa is milky-white and 0.28 mm. in
diameter. It is laid between 6 and j o'clock in the afternoon during December, in the Medi-
terranean. The segmentation is total, but not equal, and the resulting blastomeres form a
very irregularly shaped, i-layered mass, inclosing a large segmentation cavity. Finally, a
symmetrical, egg-shaped, ciliated, planula larva is formed. The planula attaches itself and
becomes a branched hydrorhiza, from which the spindle-shaped clava-like hydranths arise.
These develop in about 18 days after the egg is laid. The polypite has 13 tentacles arranged
alternately in 3 whorls.

In its irregularly shaped larva and in its mode of attachment this medusa resembles
Turritopsis, to which indeed it is evidently closely related, the only distinction between
Turritopsis and Oceania being that in the former the peduncle is vacuolated and in Oceania
it is solid and gelatinous.

.\\THO\IUH s ]•: STYI.AI TIS.

1 10

Genus STYLACTIS Allman, 1864.

Slylaclis, ALLMAN, 1864, Annals anil Mag. Nat. Hist., scr. 3, vol. 13, p. 353; 1871, Monograph TubuUnan Hy>lroi,l;,, p. 301.
CLARKF., 1882, Mem. Boston Sue. Nat. Hist., vol. 3, p. IjS. SH.IKHIOS, iSSi), American Naturalist, vol. 33, p. 801.

These are Margelinae in which the medusa is degenerate, and indeed in the European
species of Stylactis no medusae are set free, mere sporosacs being developed on the hydranths.
The two American forms, however, produce free-swimming, although degenerate medusae.

The medusa has 8 rudimentary tentacle-bulbs, 4 radial and 4 interradial ; 4 simple radial-
canals and a ring-canal. The manuhiium is sac-like without a mouth, and without oral
tentacles. The gonad encircles the sac-like manubrium, and the mature eggs or spurn arc-
found in the ectoderm. The sexes are separate.

The hydroid, Stylactis, hears a resemblance to hoth HyJractinia and Podocoryne, hut
unlike the two last named the hydrorhi/.a is not covered with a fleshy layer of cu-nosarc

c

Fl<i. 8|. — Oceania urtnuta, from life, liv tlie author. Viplr /.ML,!,,,-]! .il Station, December |S, H/O-.
A. Mature female.

M. Basal part of tentacle showing ectodermal ocelli^ on its velar sule.
C. Part <>f lip, '<lin\\m^ m\\ of nettling-warts.

(hydrocaulus). The hydrorhiza of Stylactis is formed of a network of anastomosing stolons
invested with a chitinOUS pensarc, hut are without a superficial layer of naked ccenosarc.
The hydranths are simple, claviform, or cylindrical, with one or two circles ot tilitorni ten-
tacles around the oral /one. The hydranths arise singly from the hydrorhi/a and are of
two sorts, reproductive and nutritive. Spines also arise from the hydrorhi/a. The spor-
osacs, or medusa-huds, are borne upon the sides ot the reproductive hydranths m an irregular
circlet helow the tentacles.

The type species is Styl/ictis fuciola of the Mediterranean (see Allman, 1864) which
was first described by M. Sars (1857, Nyt. Mag. Naturvid., ()), under the name Podocoryne
fuciola. Allman pointed out its peculiar characters and distinguished it from Podocoryne,

One of the most remarkable species is St\ltiftii iniimi Alcock Il8t)2, Annals and Mag.
Nat. Hist., ser. 6, vol. 10, p. 207, I fig.). This is commensal upon a fish Minum in,-rtnn.
Sporosacs, but no medusa-buds, are produced by the hydroid ot this species.

].r)0

MEDUS.E OF THE WOULD.

Stylactis hooped Sigerfoos.

Xnlii'tn hooperi, SIUERFOOS, 1899, American Naturalist, vol. 33, p. 801, 5 figs.— NuT-nun, 1901, Bull. U. S. Fish Commission
fur iS.j9, vol. ii), pp. 335, 374, figs. 13, 85. — HAROITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 41.

Medusa. — The medusa is degenerate and intermediate in its character between the
sessile sporosacs of H \Jrtictinia and the well-developed, free-swimming medusae of Podo-
CDI via'. The medusae of Stylactis hooper I live only a few hours after being set free from the
hydroid, and die soon after setting free their genital products. The sexes are separate, the
mature genital products being developed within a large part of the ectoderm of the manu-
brium, so that the gonad encircles it as in the Codonidae. The evident relationship of the
hydroid to PoJocoryne, however, necessitates the medusa's being classified in connection with
Podocoryne,

st-

B

Fi<;. 82. — .S'/y/rtr/;s /joo/ipr/, after Sigerfoos, in American Naturalist.

A. Colony shell of living llvanassa obsolete, represented as crawling over a blade of eelgrass. Natural size.

B. Part of colony magnified 6 diameters, (i) large nutritive hydranth fully expanded ; (2) same slightly

contracted; (3) same fully contracted; (4) large reproductive hydranth fully expanded; (5) smaller
reproductive hydranth, slightly contracted ; hr., tubes of hydrorhi/a ; sp., spines.

C. Recently liberated female medusa, before extrusion of eggs, cc., circular canal; re., radial-canal; s/.,

Mnrnach; ;., tentacles ; z>., velum ; ov., ova. X 50.

The bell is somewhat killer than a hemisphere, with thin, uniform walls. There are 8
short, marginal tentacles, 4 radial and 4 interradial. The velum is well developed. There
are 4 straight, slender radial-canals and a ring-canal. The manubrium is wide and spindle-
shaped and fills the greater part of the bell-cavity. There is no mouth-opening and there
are no oral tentacles. The ectoderm around the middle region of the manubrium is much
distended with the genital products.

Hydroid. — The hydroid was found by Sigerfoos growing upon the living shells of
Ilyanassa, especially upon snails found crawling upon eelgrass. It was found in Lloyds
Harbor, Huntington Bay, in Long Island Sound, New York, during the summers of 1898 99.

ANTHOMKOrs.K STVI. UTIS, TH \ M \( ISTV I.I'S. I .") I

There is no hydrocaulus. The hydrorhiza is a network of tubes lying in one plane, from
which arise a tew small, simple spines and many sessile h\ dranrhs. The hydranths are of
two kinds, nutritive and reproductive. The nutritive hvdranths are ahout 25 mm. long and
are elongate, simple, and unbranched. Their oral ends taper gradually, being widest at the
circlet ot tentacles. There are usually about 20 long, slender tentacles arranged in a single
whorl, but specimens are found with as many as 35 and as few as 15 tentacles. The entoder-
mal cores of these tentacles are solid and consist of a single row of cells. The tips of these
tentacles are not appreciably swollen or knob-like, although they are well provided with
nematocysts.

The reproductive hydranths are slightly smaller than the nutritive ones. They have
only 6 to 10 tentacles and are only one-third to three-fourths as long as the nutritive hydranths,
but they have functional mouths. 1 here are usually 4 to 5 medusa-buds arranged in a circlet
immediately below the circle of tentacles. The entoderm of the hydroid is white or pinkish-
white. The medusae appear to be set free soon after dark, and the breeding season extends
throughout July and August.

There are no spiral protective zooids such as are found in Hydractinia and there is no
coenosarc layer over the hydrorhiza such as is found in H \Jrtu tnnn and l)'iJ(n-r>r\>if.

Stylactis arge Clarke.
Xijiiilis argt, CHRKK, |S82, Mem. Boston Soc. N.u. Hi^t., \.,l. ;, p. i ^K, plate 8, figs. 18-20.

The medusiform gonophores of this species often become mature while attached to the
hydroid stock. In other cases, however, the gonophores become free, although they are
sexually mature at the time of liberation.

hit-f medusa. — The bell is ellipsoidal and the gelatinous substance thin. There are 8
small rudimentary tentacle-bulbs, 4 radial and 4 interradial. 4 simple, narrow radial-canals.
1 he manubnum is globular and greatly distended by the contained genital products.

Hydroid. — The hydranths are very much elongated and slender. Kach Indranth is
provided with 10 to 30 tentacles which arise in 2 circlets close together below the lupostome.
The tentacles ot the lower circlet are somewhat shorter than those of the upper one, although
in some individuals all ot the tentacles are of equal length. There are usually 4 sporosacs or
medusa-buds developed upon the sides ot the body ot the hydranth beneath the circlets of
tentacles, 2 being first developed on opposite sides ot the hydranth, and then 2 others appear
at points 90° from the first pair. The entire colony is of a delicate opaque white.

It is remarkable that in this species a constriction sometimes appears around the hodv
ot a hydranth, from a point just above which two or three cylindrical, stolon-like processes
are developed. The constriction then becomes complete and the distal end ot the hydranth
becomes tree. This then settles down in a new locality and the stolon-like processes give
rise to a hydrorhiza and a new colony is thus developed asexually.

This species has been found by Clarke in Chesapeake Bay.

Genus THAMNOSTYLUS.

Thardnostylut, H\K KM., 1X74, Syst. »lrr iUfilusrn, p. S^; iSSl, Rrpnrt H. M. S. ('.ltlill-'n ^ •> • . '/.•»'}., \ol. 4, p. z.

The only known species is Thamnostylui Jnh'nni llaeckel, from the Antarctic Ocean.

GKNKRIC CHARAC I I RS.

Margelinz with branched oral tentacles and 2 marginal tentacles iSo" apart. Develop-
ment unknown.

Thamnostylus dinema Haeckel.

Thamnostylui <ltnetmi, HAH KM, 1^7*), S\st. *!i-r Mrdusrn, p.N>; iXSi, Report H. M. S. (:imll>-ny, /.o<,l., vnl. 4, p. 2, pl.itr i,
8 figs.

Bell hemispherical, ]<i mm. wide, S mm. high, with thick gelatinous substance, but
thin at margin. The margin is thickly studded with nematocvsts and the exumbrella is .I!M>
besprinkled with isolated nettling cells. 2 diametrically opposite, radiall) situated, hollow.

152 MEDUS.E OF THK WOULD.

marginal tentacles about 3 times as long as the bell-diameter and ringed at frequent inter-
vals with swollen masses oi" nematocysts. The basal bulbs of these tentacles are swollen
and conical and each one bears a dark-red abaxial ocellus. There are 2 rudimentary, ocellated
tentacle-bulbs upon the margin 90° apart from the long tentacles. Velum wide. 4 straight,
narrow radial-canals and a simple ring-canal.

The stomach is a wide, 4-sided pyramid, as long as the depth of the bell-cavity. It extends
below into a 4-sided, elongated throat-tube, twice as long as the stomach itself. This throat-
tube projects beyond the velar opening and the mouth is a simple, 4-sided opening studded
with nematocysts.

4 large, radially placed, oral tentacles arise from the distal end of the stomach and branch
dichotomously 6 to 8 times, their ends terminating in nematocyst-knobs. The gonads are
swollen, leaf-like, lobular, folded ridges in the 8 adradii of the stomach. They are blunt
above, but more pointed below, and each gonad exhibits about 5 double transverse folds.
The ova are prominent. The gonads, oral tentacles, nematocysts of bell-margin and of the
tentacles and tentacle-bulbs are blood-red.

Antarctic Ocean, south of the Kerguelen Islands, at a depth of 120 fathoms. Beautifully
figured by Haeckel, 1881.

Genus THAMNITIS Haeckel, 1879.

Thamnitis, HAF.CKEL, 1879, Syst. der Medusen, p. 84. — VANHUFFF.N, 1891, Zool. Anzeigcr, Bd. 14, p. 444. — BROWNE, 1905, Proc.
Royal Soc. Edinburgh, vol. 25, p. 758.

The first described species of this genus is Thamnitis nigritella, first made known by
Forbes, 1848, under the name Bougainvillea nigritella. The first species mentioned by
Haeckel is a newly-described form, Thamnitis tctrclla. T. nigritella comes from the Shetland
Islands, while T . tctrclla was obtained off the coast of Brazil.

GENERIC CHARACTERS.

Margelinae with branched oral tentacles and 4 simple, radially placed, marginal tentacles.

It is possible that Thamnitis may be simply an immature or damaged Bougainvillia
which has lost all but one marginal tentacle of each radial cluster, but we are not yet justified
in concluding that this is the case. Forbes, 1848, describes and figures the tentacle-bulbs of
Thamnitis nigritella as being bean-shaped, containing a single large ocellus ( ?) or mass of
very dark pigment, and having a single short tentacle arising from one side of the bulb, not
from the middle. Haeckel, 1879, p. 84, describes T. tctrclla as having small, pear-shaped ten-
tacle-bulbs with a single tentacle arising from the middle of each bulb. He does not mention
ocelli, but his description is brief and unaccompanied by a figure. Further study is required
before we can be certain that T hamnitis actually exists in nature. Vanhoffen, 1891, accepts
it with some hesitation, and I share his view in this respect.

Browne, 1905 (Proc. Roy. Soc. Edinburgh, vol. 25, p. 758), is the only recent author
who has seen a specimen of Thamnitis. He found two small, immature medusae in the Firth
of Clyde, Scotland, in September to October. They were I to 1.5 mm. wide and had 4 simple,
marginal tentacles. No ocelli. 4 oral tentacles 2 to 3 times dichotomously branched. No
gonads. Stomach yellow to orange, tentacle-bulbs orange.

"Thamnitis nigritella" Forbes, 1848 (British Naked-eyed Medusae, p. 63, plate 12,
fig. 2) is probably an immature Bongainvillia which has lost all but one tentacle in each
marginal cluster.

Thamnitis tetrella Haeckel.

Thamnilii lelrel/a, HAECKF.L, 1879, Syst. der Medusen, p. 84.

Bell egg-shaped, being 6 mm. in height and 4 mm. in diameter. There are 4 radially
situated, marginal tentacles which are longer than the bell-height. Each tentacle arises from
the middle of a large basal bulb. An ocellus is situated at the base of each tentacle. There
are 4 radial-canals. The manubrium is cubical with a wide quadratic base. There are 4
radially situated, oral tentacles, each of which branches dichotomously three times. The
gonads are found in 4 interradial ( ?) swellings upon the wall of the stomach. Color ( ?)
This species is found upon the coast of Brazil.

ANTHOMKWS.K I.V\1\( HtKA. 1~>:!

Genus LYMNOREA P«ron et Lesueur.

( ?) Fnfonia, PERON FT LCSIIKUR, 1809, Hist. Gen. des Meduses, p. 16.

Lymnorea, PERON KT LESUEI'R, 1809, Ann. Mus. Hist. Nat. Paris, tome 14, p. 329. — DE BHINVIM i, 1834, Manuel il'Actim.l.,

p. 290. — MAYER, 1900, Bull. Mus. Cmnp. Zool. at Har\ard College, vol. ',7, p. 6; 1904, Mem. .N.n. Su. Brooklyn Inst.

Museum, vol. i, p. io.— Ar.ASStz and MAVER, 1902, Mrm. Mus. Gimp. Zool. at Harvard College, v..l. i(,. p. 144.
Limnorra+ Thaninmtonui, HAKTKM-, 1879, Syst. der Medusen, pp. 85, 86.
l.tmnurfu^- Thomiiiutoma, VANHOFFEK, 1891, Zool. An/eiger, Bd. 14, p. 445.
MiHtadia, BROOKS and RlTTENHOUSE, 1907, Proc. Boston Soc. Nat. Hist., vol. 33, p. 437.

The oldest species of this genus is l.\tnnur,-a tri,;im, described by Peron and Lesueur,
and first figured by de Blainville, 1834. It occurs on the southern coast of Australia. The
figures and description leave much to be desired, but the branching oral tentacles are clearh
represented. Tht- marginal tentacles are small and numerous.

GENERIC CHARACTERS.

Margelinx with branched oral tentacles and with 8 or more marginal tentacles which
arise separately, not in clusters, from the bell-margin. The hydroid is unknown.

Haeckel, 1879, would call medusae with 8 marginal tentacles Thamnostoma, and those
with 16 or more Lnnnori-u. He introduced the spelling Limnorca, but Peron and Lesueur
spell it Lymnorea. Owing to the very incomplete description of Peron and Lesueur it is
impossible to determine whether Favonia is or is not equivalent to Lymnorea. Judging from
de Blamville's, 1834, copy ot Peron and Lesueur's unpublished figure, it appears to be closely
allied to Lymnorea, but its relationships must remain uncertain, l-'tivnnia takes precedence
over Lymnorea, should it be equivalent, for it precedes Lymnorea in Peron and Lesueur's
description.

Lymnorea dibalia.

Liszia dibalia, BUSCH, iSijl, Beohach. wirbellos. Seeth., p. 23, taf. 1, fign. 7-9.

Ttiumnostomti dibaJis, HAEOKEL, 1879, Svst. der Medusen, p. 86. — GRAEFKE, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 348.

Hell oval, 7 mm. high, 6 mm. wide. 8 marginal tentacles, those at the bases of the 4
radial-canals being longer than the 4 interradial tentacles. The ocelli are borne upon pro-
jections, one ot which arises from each tentacle-bulb. The stomach is cubical and lacks a
peduncle, and the throat-tube is as long as the stomach. There are 4 oral tentacles which
branch dichotomously twice. 4 swollen gonads on the interradial sides of the stomach.

Found at Trieste, Adriatic Sea, Mediterranean. Graefte states that the mature medusje
are found in October, and the young first appear in September.

Lymnorea macrostoma.

Thamnostoma rnacro<.tomti, HAK'KH , iSy9, Svst. der Medusen, p. 86.

Hell cubical, 8 mm. high, 8 mm. wide. 8 similar marginal tentacles, each longer than the
bell-diameter. A simple, abaxial ocellus at tin- base ot each tentacle. Stomach globular with-
out a peduncle. The throat-tube is 3 times as long as the stomach and extends beyond tin-
velar opening. There are 4 short, oral tentacles, each of which branches dichotomously o to 8
times, thus giving rise to 80 to IOO terminal branches in each (jiiadrant. 4 gonads on sides of
Stomach. Color ( ?) Indian Ocean, Singapore.

Lymnorea ocellata Agassiz and Mayer.

Lymaorea afdlntti, At,\ssi/., A., and MAYER, 1902, Mem. Museum Comp.Zool.al Harvard CoU*i vol. 16, p. 1441 plate 2, figs. 9-11,

( ') I.ymnorfa Iritdra, Pi KON tr I.tsi n R, 1X09, Anna), du Mus. Hist. Nat. P. ins, tome 14, p. 521); dc- Bl.niu ille, lH)4, Man.

Actinolnt-H-, p. 2(jo, plate 40, fig. 2; HAH KM, lS79, Svst. der MrditM-n, p. 87.
( •) l.imrtorea probosciitfa, HAFCKM, iS?<j, Svst. der Medusen, |<. Sy.
( f)Limnorea probosfiilta, BRDOT, 1901, Revue Suisse de Zoo!., tome 9, p. 4X4; Ibitl., 1905, tttine i',, p. 137.

The top ot the bell is flat and the sides flare outward in a bell-shaped manner. The bell
is about 4 mm. in diameter and is about as high as it is broad. The walls are thin and quite
flexible. There are about 50 short, stiff" tentacles which are each about half the length of the
bell-height. These tentacles are usually carried curled upward and their entodermal cores are
solid and composed of vacuolatcd chordate cells. The basal bulbs of the tentacles are large

I.-, I

MF.m'S.E OF THE WORLD.

Fir.. 8}.--/, \mnorea offlliita, after Agassiz and
Mayer, in Mem. Museum Comp.
Zool. at Harvard College.

and each contains a mass of red entodermal pigment. In addition to this there is a prominent
ectodermal ocellus upon the velar side of each tentacle at a short distance outward from the
basal bulb. The velum is narrow. There are 4 straight, narrow radial-canals and a slender cir-
cular vessel. The peduncle of the manubrium is wide and quadratic in cross-section and its
lower portion near the stomach consists of highly vacuolated, entodermal cells. The stomach is
pear-shaped and the mouth is a simple, round opening. The entire manubrium extends about

half the distance from the inner apex of the bell-cavity
to the level of the velar opening. There are 4 well-
developed, radially placed, oral tentacles, each one of
which is about half as long as the height of the manu-
brium. Each oral tentacle branches dichotomously 4
times, thus giving rise to 16 distal knobs. Each knob is
thicklycovered with a bristling cluster of fusiform nema-
tocyst-cells. In addition to these there are several
patches of nematocystic bristles upon the sides of the
main shaft of each oral tentacle. The gonads are devel-
oped in 4 interradial regions on the walls of the stomach.
The entoderm of the tentacle-bulbs and of the stomach
is brick-red. The ectodermal ocelli are black. A num-
ber of these medusae were obtained on the surface in
the lagoon of Makemo Island, Paumotus, South Pacific, on October 23, 1899.

This species may be identical with Lymnorea triedra Peron et Lesueur = L. probosciJea
Haeckel, but the published drawing of this medusa is evidently inaccurate and the description
so brief and vague that it will probably never be possible to redetermine the species. /,. trn'Jrn
is described from Bass Strait, between Australia and Tasmania.

Lymnorea borealis Mayer.
Plate 15, figs. I to 3.

Lvmnorea borealis, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. (>, figs. 16-18, plate 5.

( ;} Limnorea tioivegica, BROCH, 1905, Bergens Museums Aarbog, No. 11, p. 5.

(?) Cvttftinda nreolala, BROWNF., 1897, Proc. Zool. Soc. London, p. 817, plate 48, figs. I, 2. (Podocorvtif!')

Bell 3 mm. high. It is higher than a hemisphere and is acorn-shaped, with a low, blunt
apical projection. The bell-walls are thin. There are 32 marginal tentacles, each about halt
as long as the bell-height. They are quite stiffand are carried curled upward, above the margin.
Their basal bulbs are large and densely pigmented. The velum is well developed. There are
4 straight, narrow radial-canals and a slender, simple, circular vessel. The manubrium is
slender and pyriform and mounted upon a very short, solid peduncle. It is cruciform in cross-
section. The mouth is flanked on its 4 radial corners by 4 short dichotomously branching oral
tentacles (plate 15, fig. 2). Each oral tentacle branches dichotomously twice, thus giving 4
tentacle tips in each quadrant. These tentacle tips are knob-like and armed with long spindle-
shaped nematocysts (plate 15, fig. 3) very much as are the oral tentacles of Lymnorea ocellata.
The 4 gonads are found in four longitudinal, interradial, swollen regions in the ectoderm of the
stomach-wall. The immature eggs are transparent and give a reticulate appearance to the
surface of each gonad. The entoderm of the manubrium and tentacle-bulbs is bright-red.
There are no ocelli.

Three specimens were taken on the surface at Eastport, Maine, on September 19, 1898.

Broch, 1905, gives a brief description of a Lymnorea from the coast of Norway which may
be identical with this species. He states that there is a pair of very short, oral tentacles at each
of the 4 corners of the mouth and that each of these forks dichotomously twice, thus giving 8
terminal knobs at each radial corner of the mouth. There are 16 to 22 tentacles. In other
respects his description accords with that of/,, borealis, allowance being made for contraction
due to preservation in Broch's specimen. He gives no figures and does not mention the color.

Lymnorea alexandri Mayer.
Plate 15, figs. 4 to 9.

Lvmnorea ale\andri, MAVF.R, 1906, Mem. Nat. Scj. Museum Brooklyn Institute, vol. i, p. 10, plate I, figs. 1-5*7.
M.ir.i.lm, i/)., BROOKS an. I KITTKNHOHSF., 1907, Proc. Boston Soc. Nat. Hist., vol. 33, p. 435, plate 31, fig. 8.

ANTHOMEDUSjE— r.VMXoltr.A, BOUQAINVILLIA.

Bell 4 mm. high, 3 mm. broad, with vertical sides, flat top, and thin walls. 32 slender ten-
tacles with tightly coiled ends and large basal bulbs. A black, ectodermal ocellus lies upon the
velar side ot each tentacle-bulb ( plate I 5, fig. g). The velum is well developed. The 4 radial-
canals are straight and slender, but slightly swollen in the mid-region of their lengths, where
their inner sides are lined with gland-cells (plate i 5, fig. 6). The manubrium is flask-shaped,
and about halt as long as the depth ot the bell-cavity. The distal part of the manubrium ad-
jacent to the points ot entrance ot the 4 radial-canals is composed ot highly vacuolated cells.
The mid-region ot the manubrium is cruciform in cross-section and the mature genital pro-
ducts are found in the ectoderm ot the 4 interradial sides of the stomach. The mouth is at the
extremity ot a short neck and is surrounded by 4 oral tentacles, each ot which branches dichot-
omously ^ rimes and terminates in 8 nematocvst-beanng knobs, making in all 32 ot these
knobs surrounding the mouth (plate i 5, fig. 5 ). The entoderm of the manubrium and tentacle-
bulbs is dull, flesh-colored pink. In the youngest medusa seen the bell was O.(> mm. in diameter
and had only 4 oral tentacle-knobs, 4 marginal tentacles, and 4 rudimentary tentacle-bulbs.
This is one ot the most abundant medusae upon the surface in summer, among the Bahama
Islands. A single specimen was found at Tortugas, Florida, in July, igo5. Named in honor
of Dr. Alexander Agassiz.

Genus BOUGAINVILLIA Lesson, 1836.

Cvan<ra bou^ainvillii, LESSON, 1830, Voyage de la Cor/ui/le, Zool., torn, z, part. 2, ir Jiv., p. I 18, plate 14, Zoophytes, figs. 3,

D,D',D",D'".

Bougainvillia, LKSSON, 1836, Ann. ties Sci. Nat., Zool., scr. 2, torn. 5, p. 262.
Hi/>[>o(rene, BRANDT, (Mertens), 1835, Mem. Acad. Imperial)- des Sci. St. Petersliourg, Sci. Nat., SIT. 6, tome 2, p.22i). -Pre-

occupied by OKEN, 1X17, for Mollusca.- -Ac.ASSiz, L., 1849, Mem. Amer. Acail., NYu Strie--, vol. 4, p. 250.
Bougainvilka, FORBES, 1848, British Naked-eyed Medusie, p. 61.
Boiigtiinnllm, AC.ASSIZ, L., 1862, Cont. Nat. Hist. I'. S., vol. 4, p. 344.-
Lizusa (young medusa), HAECKEL, 1879, Syst. der Medusen, p. 80.
llippocrene, HAECKEL, 1879, Syst. der Medusen, p. 90.

Z, A.,

.^, North Amer. Acal., p. 152.

ppocrene, AECKEL, 179, ys. er eusen, p. 90.

BougainviUiaj HARTLAI;B, 1897, Wissen. Meeresuntersuch. auf Helgoland, Heft i, Abt. 2, p. 455.
Bougainmllea, MAAS, 1905, Ciaspedoten Medusen der Siboga Expedition, Monog. 10, p. 10.

neu i, not. 2, pp. 472, 473.

Boueainviffia, HART-IIT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 37.
Boii^ainrillfti, GI.RD, 1892, Zool. An/eiger, Bd. 15, p. 312, 5 fign. (cell-division] development of morulu).

GENERIC CHARACTERS.

Margelina; with 4 radially placed clusters
of marginal tentacles, the tentacles ot each
cluster being all of one kind and similar in
structure each to each. Hydroid: Bougain-

There are 4 radially placed, dichoto-
mously branching, oral tentacles, a quadratic
manubrium, 4 radial-canals, and gonads mi
the interradial or adradial sides ot the stom-
ach as in other Bougamvillidi.

The type species ot the genus is Bougain-
villia nmclwiiiHii Lesson, 1836, from the Falk-
land Islands. This medusa was first described
and figured by Lesson, in 1830, under the
nzmtCyaneabougainvilhi. 1 he generic name
" (1\iim-n" was, however, previously used by
Peron and Lesueur, i8og, for Discomedusa;.
\lso the name Hippocrene of Brandt (Mer-
tens), 1835, was preoccupied in iSl/byOken

for Mollusca and can not be applied to Medusa.-. It therefore becomes necessary to adopt

the generic name Bougainvillia.

fit;. 84.— Bougainvillia "museui," after Allman, in R.iv
Sin-iety, 1871-1872.

lflfl

MEDUS.E OF THE WORLD.

The name "Margelis" was first proposed by Steenstrup, 1850, and applied to designate
Margrlis principis of the North Sea, and this name was adopted hy Agassiz and Haeckel.
They would designate "Margelis" as being similar to Bougainvillia, but with a manubrium
having a narrow base devoid of a peduncle and with the 4 radial-canals opening close together
into the 4 corners of the narrow stomach. Bougainvillia, on the other hand, they would
restrict to designate medusae with a wide quadratic stomach and often with a peduncle, the
4 radial-canals arising far apart from the 4 corners of the wide cruciform stomach. While
these distinctions serve admirably to distinguish "Margelis" principis or "M." carolinensis
from Bougainvillia superciliaris, they fail completely to classify medusae in which the manu-
brium is of moderate width, neither wide nor narrow. It is my object to retain only positive.

Fir,. 85. — Bougainvillia "/r»//Yo'/j," after Allman, in Ray Society, 1871-72.

not relative, characters as of generic value. It is futile and confusing to attempt to separate
genera upon such intergrading differences as "wide or narrow stomach," "long or short manu-
brium," etc., for we generally find that species will be discovered allied evidently to both these
so-called "genera," but strictly intermediate in essential characters, Bong/iinvillia takes
precedence over Mar gel is and should therefore supplant it.

Forbes, 1848, introduced the spelling Bougainvillea, but Lesson's original proposition is
Bougainvillia.

Haeckel, 1879, p. 90, confuses the hydroid of Bougainvillia with EuJenJrium, but in
Eudendnum no free medusx are produced and there are other distinctions separating it from
Bougainvillia.

ANTHOMEDUSjE — BOUGAINV1LLJA.

157

Nemopsis is separated from Bougainvillia by the tentacles, composing its 4 radially
situated, marginal clusters, being of two distinct kinds (clubbed and simple-ended). The mar-
ginal tentacles ot Bougainvillia are all of one sort and do not terminate in knobs. Vanhoffen,
1891, shows that we can not separate Nemopsis from Bougainvillia upon the mere length of the
radial pouches of its stomach, for some medusx of Bougainvillia display pouches of consider-
able length.

Tabular Synopsis uj the Medusa of Bougainvillia.

B. macloviana
Lesson.

B.principis=

Margelis
principis
Steenstrup.

B. fulva
Agassizand
Mayer.

B. britannica
Forbes.

B. Miprrciliaris
L.Agassiz.

B. dim-ma
Greene

= B. super-
ciliaris ( r)*

B.multicilia =
Lizusa
multicilia

Haeckel.f

Shape and size

Cylindrical,

Nearly spheri-

Cylindrical,

High, dome-

Dome-like, with

Dome-like,

0.75 spherical.

of bell in

with round-

cal. 7 wide,

with flatly

like. Thick-

bulging

with bulging

6 high, 5

mm.

ed top. i 5

6 high.

rounded

walled.

sides. 10

sides. 12

wide.

high, 1 3

top. 14

high, 10

high, 8

Thin uni-

wide.

high, i i

wide .

wide.

form bell-

wide.

walls.

Number of

35 to 65 in

25 to 30 in sin-

I5to2oin sin-

1 6 to 20. Very

10 to 15.

10 to 12 ( ?)

to to 12

tentacles

double row.

gle row.

gle row.

short, in

~\2 in the

( >nlv 2 per-

upon each

single row.

variety

fect tenta-

of the 4 mar-

B. aun-j.

cles seen.

ginal bulbs.

Shape of mar-

V-shaped.

Long, linear;

Epaulet-

Epaulet-

Elongate, oval

Kidnev-

Kidney-

ginal bulbs.

wider than

shaped.

shaped.

basal bulbs,

shaped.

shaped.

spaces be-

of small size.

tween them.

Shape and

Small, cubical.

Short, wide.

Wide. Half

4-sided, \\ tile,

Wide, 4-sided,

Small, balloon-

Globular to

size of man-

On broad,

4- sided .

as long as

urn-shaped.

and flask-

shaped,

flask-shaped

ubrium.

conical

Only one-

depth of

No pedun-

shaped.

4-sided,

with con-

peduncle.

third as long

bell-cavity.

cle. Only

Mounted on

hardly one-

stricted

as depth of

No pedun-

about half

peduncle i .5

fourth as

base. \->

bell-cavity.

cle.

as long as

to two-thirds

long as bell-

peduncle.

depth of

as long as

height.

bell-cavity.

depth of bell-

cavity.

Number of

6 to 7 times.

4to5times.

8 times.

6 to 7 times.

About ^ times.

4 to 5 times.

Simple, un-

dichotomous

7 times in

branched.

branchings

the variety

of each of

B. aurea-

the 4 oral

tentacles.

Character of

4 wide, folded,

4 spherical,

8 adradial.

8 adradial.

4 interradial.

4 interradial.

8 adradial.

gonads.

interradial

interradial.

No planulac

Planulae

swellings.

remain

develop in

attached to

capsules

stomach.

upon siil'"."!"

stomach.

Color.

Stomach and

?

Stomach and

Stomach and

Stomach and

StMMi.n h and

Stomach light-

tentacle-

tentacle-

tentacle-

tentacle-

tentacle-

vello-A .

bulbs yel-

bulbs

bulbs

bulbs rosin-

bulbs yel-

Tentacle-

low. Gon-

orange, v--l-

golden-yel-

yellow to

low. Ocelli

bulbs

ads and

low, brown-

low. <>ceiii

reddish-

black.

orango

ocelli

ish-yellow.

black.

brown. * >(. el-

vell'iw.

brownish-

Ocelli

li black.

OcelH dark-

black.

bUK.

red.

Where found.

Falkland

North Sea,

Tropical

North Atlantic

Arm* ( kean,

East oust nf

Sir. nts of

Islands,

coast of

Pacific,

coasts of

northern

Ireland.

Gibraltar.

South At-

Scotland.

Fiji to

Europe and

coasts of

lantic.

Malay

America.

Europe and

Archipelago.

America.

Hydruid.

Unknown.

Unknown.

Unknown.

Hvdroid has

B. gtiperciliaris!

been com-

monly * . til-

ed B." ra-

mosa."

*This medusa appears to be B. supfreiliaris which has lost some trnt.ulrs. H. <m>r,i, I.inko, 1^04, is pmbablv a \.irii-iv <-\
B. Mtpcrciliarh from Arctic Ocean.
fAn immature medusa?

l.-.s

MEUUS.K OF THK WOULD.

The definitions of Bougainvillia and Nemopsis here proposed are identical with those of
Hargitt, 1904, p. 37.

Species of Bougainvillia are widely distributed in all seas, but are especially abundant in
the cold waters of the northern hemisphere, a remarkably large number being found in the
North Sea near Helgoland, by Hartlaub, 1897. Many of the species are very closely related
and their distinctions may be due in part to environmental influences. Hallez found that

"T ulndar Synopsis of the Medusa nj Bougainvillia. — Continued.

H.pl.itygaster
Haeckel.

B. xantha
Hartlaub.*

B. carolinen-
sis= Hip-
pocrene
carolinensis
McCrady.

B. niobe
Mayer.

B.pyramidata
= Hippo-
crene pyra-
midata
Forbes.

B. nordgadrdi
= Margelis
nordgaardii
Browne .

B. flavid.i
Hartlaub.

Shape and size

Cubical.

Dome-like,

Dome-like.

Cylindrical,

Semi-globular.

Oval. 4 high,

Oval. 3 wide,

of bell in

12 high, 12

walls very

Very thick

with flatly-

6 high, 6 wide

4 wide.

2 high.

mm.

wide.

thick. 6

walls. 4

rounded

Walls thick.

Walls only

Walls thick.

high, 5 wide

high, 4 wide

apex. 6.75

fairly thick.

high, 4.8

wide. Bell-

walls thick.

Number of

10 to 12

9 to to

7 to 9 very

8

6 to 8

5 toy. No

Not more than

tentacles

long and

ocelli.

6.

upon each

slender.

uf the 4 mar-

ginal bulbs.

Shape of mar-

Triangular,

Small, oval to

Small, thick,

Small, oval.

Small, globular.

Small, globu-

Swollen, fairly

ginal bulbs.

wider than

globular.

bulbous.

lar.

large, bul-

spaces be-

bous.

tween them.

Shape and size

Flat, 4-sided.

Spindle-shap-

Narrow,

Flask-shaped.

Flat, cruciform,

Half aslong as

Small, conical

of manu-

No pedun-

ed. No ped-

widest near

Cruciform

mounted

depth of

about h.ilf

brium.

cle.

uncle.

mouth.

in cross-

upon pedun- ' bell-cavity.

as long as

Half as long

section .

cle.

depth of

as depth of

Half as long

bell-cavity.

bell-cavity.

as depth of

No pedun-

bell-cavity.

cle.

Number of di-

3 or 4 time&.

About 2 or 3

2 times.

4 times.

2 times.

4 or 5 times.

3 times.

chotomous

times.

branchings

of each of

the 4 oral

tentacles.

Character of

4 interradial.

4 interradial.

4 interradial.

Medusa-buds

GonaQE itn base

4 interradial.

4 interradial.

gonads.

arise from 8

and lobes of

adradial

stomach.

sides of

stomach.

Color.

Stomach and

Stomach and

Variable, us-

Stomach and

Stomach and

Gonads and

Stomach and

tentacle-

tentacle-

ually stom-

tentacle-

tentacle-

tentacle-

tentacle-

bulbs red-

bulbs gold-

ach and

bulbs rosin-

bulbs orange

bulbs yel-

bulbs

dish-yellow.

en-yellow,

tentacle-

yellow.

to yellow to

lowish-

orange-yel-

Ocelli ?

reddish-yel-

bulbs green

reddish-

brown (in

low. Ocelli

low. Ocelli

or green and

brown.

formalin).

black.

small,

red. Ocelli

No ocelli.

brownish-

black.

violet.

Where found.

Eastern parts

Helgoland,

Atlantic coast

Bali am a

West coast of

Near Bergen,

Helgoland,

of tropical

German

of United

Islands,

Scotland,

Norway-

German

Atlantic.

Ocean.

States,

tropical

south of

Ocean,

Cape Vrnlr,

Woods

Atlantic.

Ireland.

August to

Canary

Hole to

September.

Islands.

Tortugas.

Hydroid.

Unknown.

Unknown.

Hydroid: B.

Unknown.

Unknown.

Unknown.

Hydroid: B.

carolinensi.s.

ffavida

Hartlaub.

*Medusa allied to B. fariJa

AXTHiiMKDI s.K inn c \L\V1I.I.IA.

159

B. frutic'jsa Allman is a form of B. ramosa Van Bencden, which grows in agitated water.
The ohservations of GraefFe, 1884 (Arbeit. Zool. Inst. Wien., Bel. 5, p. 34()), are interesting in
the same connection, for he found that five distinct forms ot Bougainvillia medusx arise from
a hydroid apparently identical with Bougainvillia muscus Allman. It is therefore prohahle
that many of our so-called "species" of Bou«nim'illia are merely environmental races. The
hydroids of most of them remain unknown, or are not associated with any known medusa,
and many of the species are imperfectly described. The following descriptions must he con-
sidered as only tentative attempts to unravel the confusion which prevails at present and
which can not be dissipated until the hydroids of all or most of the species are discm 1 1< d.
Haeckel's 1879, genus " LizHsii" appears to apply only to immature specimens of lion-
gainvillia in which the oral tentacles have not \ et become branched.

Tabuliir Synopsis nj the Medusa ») Bougainvillia.* — Continued.

B.autumnalis
Hartlaub = B.
gibbsi Mayer.

B. prolifera —

Luusa prolif-
era von Len-
denfcld.

B. maniculata
Haeckel.

B. rut;o ,i
Clark.-.

B. trinema =
M jrgelis tri-
ne ma von
Lcndenfeld.

B. fnmdosa
Mayer.

.Shape and size

Dome-like.

Dome-like.

Nrjrlv spherical.

Pyriform.

Oval.

Dome-like with

of bell in mm. £.5 high, 2

3 high, 2. 5

i.i; high, 1-5 1-5 high.

3 widr.

vertical sides.

wide.

wide.

wide.

2 high, i .2^

Thick walls.

Thin walls.

wide. \\..ll

thick.

Number of ten-

3 to 6

5

4 short, finger- 3

3

2. No ocelli.

tacles upon

shaped.

each of the 4

marginal

bulbs.

Shape of mar- Rounded, bul-

K.ugc, " scrota-

Small, globular. Small, globular. Small.

Small, w ithout

ginal bulbs. bous,fairlv

shaped."

ocelli.

large.

Shape and size Short urn-shap-

Small, nearly Flask-shaped. Short, thick,

Small.

Cruciform, short.

of manubnum. ed, wide, 4-

cubical.

Narrow base, flask-shaped.

thick, flabk-

sided. Not

spherical

shaped. Half

half as long as

stomach.

as long as depth

depth of bell-

Long, conical

of beil-cavitv .

cavity. No

throat.

peduncle.

Number of di- 2 or 3 times. Simple un-

Twice. Onlv Unbranched, Kai.li end* m ;

2 or 3 times.

chotomous

branched.

tips branched. simple. branch'-:-.

branchings of

each of the 4

oral tentacle^.

Character of

4 interradial.

Medusa-buds on

4 interradial. Not developed

? Not developed

8 adradial. Plan-

gonads.

sides of stom-

(immature).

ula- develop

ach.

upon M !•• of

stomach.

Color.

Slum, H li grren-

Stomach .tii'l

Stomach and

Stomach and

Stomach and

Stomach an.! it n-

brown. Ten-

tentacle-bulb:-

tnii. 11 lr-bulbs

tentacle-bulbs

tentacle-bulbs

tacle-bulbs

tacle-bulbs

brown.

violet-gray.

vrllow, brown.

dark-vrlln\\ .

pinkish-white.

yellowish-

Tentacle-tips

greenish -yel-

turquoise.

low. Ocelli

black.

Where found.

Newport, United

Port Jackson,

Mediterranean.

Virginia in s, ,nth

Port Jat k ' 11,

Tortugas>.

States. Helgo-

New South

Carolina,

New South

land, (lerman

Wales March

Atlantic coast

U'aK-s, Aus-

Ocean. Aug-

to April.

of United

tralia.

ust to No\ em-

States.

• '

ber, and off

British coasts.

Hydroid.

Hydroid un-

An imm.iturr 1'nknown. (hdnud: I*

\ iiknown.

Unknown.

known.

form ? Un-

rugosa Clarke

known.

*For B. obscitru B»iini-vir and B. ghnella Torrry, see tclt.

160 MEDUS.E OF THE WORLD.

There is considerable variability in coloration among individuals of the same species in
many forms of Bougainvillia. The preceding tables present a synopsis of the so-called
"species," although it is probable that some of the small medusa listed near the end are
only the young of those described higher up on the table.

Gerd, 1892, finds that the segmentation in the egg of Bouganivillin is at first total and
equal and a central segmentation cavity is formed. Then the cells divide tangentially and
the nuclei take up a peripheral position. Later there is multipolar ingression into the segmen-
tation cavity and thus a solid morula is formed, having all cells similar each to each. The
peripheral layer of cells forms the ectoderm of the planula larva. Before the formation of the
ectoderm the cell boundaries become indistinct as in Turntopsis, but later they reestablish
themselves.

Bougainvillia macloviana Lesson.

Cyanea bougainvillii, LESSON, 1830, Voyage dc la Coquille, Zoo]., tome 2, part 2, 2d div., Zoophytes, p. 118, plate 14, figs. 3,

D-D'".

Bougainvillea macloritina, LESSON, 1843, Hist. Zooph. Acal., p. 290. — AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344.
Hippocrene macloviana, HAECKEL, 1879, Syst. der Medusen, p. 90, taf. 5, fign. I, 2. — BROWNE, 1902, Annals and Mag. Nat. Hist.,

ser. 7, vol. 9, p. 278. — BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 135 (literature 1830-50).

Adult medusa. — Umbrella a little higher than broad, about 15 mm. high, with a quad-
rangular margin, and with longitudinal furrows on the exumbrella. There are 4 V-shaped,
radially situated tentacle-bulbs, each giving rise to about 35 to 65 tentacles arranged in a
double row. There is a single ocellus at the base of every tentacle. The stomach is small, cubi-
cal, and situated upon a broad, cone-shaped peduncle. 4 narrow, perradial lobes extend out-
ward from the stomach along the whole length of the peduncle. The mouth is small, with 4
short, perradial lips placed at a point about two-thirds down the depth of the umbrella-cavity.
There are 4 closely and thickly branched tentacles, each with 80 to 100 terminal branches
ending in knob-like clusters of nematocysts. The gonads are developed upon the interradial
sides of the stomach and extend over the sides of the perradial lobes of the stomach, hang-
ing down in a folded band from the peduncle. The stomach and tentacle-bulbs are yellow,
while the gonads and ocelli are brownish to black. This species is abundant at the Falkland
Islands. Mature specimens were recently obtained by Vallentin between November and
February at Stanley Harbor and described by Browne. Haeckel's specimens appear to
have been immature.

Bougainvillia principis.

Margelis principis, STEENSTRTP, 1850, Vidensk. Meddel. Nat. For. Kjobenhavn, p. 35.

Bougainvillea fruticosa+ B. dllmani, ROMANES, 1876-77, Journal Linnean Soc. London, Zoo!., vols. 12, 13, pp. 526, 190.
Margelis principis, HAECKEL, 1879, Syst. der Medusen, p. 88, taf. 6, fign. 14-16. — BROWNE, 1895, Proc. and Trans. Liverpool
Biol. Soc., vol. 9, p. 266.

Bell nearly spherical, 6 to 7 mm. wide, 5.6 mm. high. Bell-walls only moderately thick
and quite uniform. 4 marginal clusters of tentacles. Each cluster arises from an epaulet-
shaped base, and contains 25 to 30 short tentacles, shorter than the bell-radius. There is an
ectodermal ocellus on the velar side of each tentacle near its base. The basal epaulets of the
tentacles are somewhat wider than the interradial parts of the margin between them.

The stomach is short, swollen, and without a peduncle. It is not half as long as the depth
of the bell-cavity, and its interradial sides are occupied by 4 swollen gonads. The 4 oral
tentacles each branch dichotomously 4 to 5 times, thus giving 20 to 30 knob-shaped, terminal
branches in each quadrant. Color ( ?)

Found off the eastern coast of Scotland and in the North Sea, at Far-Oer Island.

The hydroid is unknown.

This medusa is distinguished by its short stomach and the remarkable length of its mar-
ginal tentacle-epaulets, which are linear, not triangular, in outline.

Bougainvillia fulva Agassiz and Mayer.

Bougainvillea fulva, AGASSIZ, A., and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 162, plate 2, fig. (>;
Ibid., 1902, Mem. Mus. Comp. Zool. at Harvard Coll., vol. z6, p. 145, plate 2, fig. 8. — MAAS, 1905, Craspedoten
M-Musen der Sibog/i Exped., Monog. 10, p. 10, taf. I, fig. 8; taf. 2, fign. 9, 10: 1906, Revue Suisse de Zool., tomr 14,
p. Sy, plate 2, figs. 4, 5. — BIGELOW, H. B., 1909, Mrm. Mus. Comp. Zool. at Harvard College, vol. 37, p. 195, plates 6
and 44.

PLATE 17.

Fig. I. Bou%ainvillia supt-rciliaris. Showing planulae developing upon the
interradial sides of the manubrium. Woods Hole, Massachu-
setts.

Fig. 2. Bougainvillia nigosa, young medusa. Charleston Harbor, South
Carolina, December 17, 1904.

Fig. 3. Bougainvillia tuitiiinnalis, young medusa. Agassiz Laboratory,
Newport, Rhode Island, July, 1902.

Fig. 4. Bougainvillia autumnalis, mature female. Agassiz Laboratory,
Newport, Rhode Island, August, 1892.

Fig. 5. Nemopsis bachei. One of the median marginal tentacles of the
medusa shown in figure 6.

Fig. 6. 'Nemopsis baflici, mature male. Hampton Roads, Virginia,
November I, 1904.

Fig. 7. Bougainvillia carolinensis, mature male. Charleston Harbor,
South Carolina, September 10, 1897.

Fig. 8. Bougainvillia britannica, mature female. Eastport, Maine, Sep-
tember 19, 1898.

Drawn from life, by the author.

PLATE 17

.

ANTHOMEDU8.fi— BOUGAINVILLIA.

161

The mature medusa is described by Maas, Dr. Agassiz and his assistant ha\ ing found only
immature specimens.

Mature tni'Jusa. — Bell 8 to 14 mm. high, with thick, vertical sides and flatly rounded top,
4 radially placed clusters of marginal tentacles with about 15 to 20 tentacles in each cluster.
These tentacles are not half as long as the bell-height and are clavate and flexible. There is
an ocellus on the velar (inner) side of the base of each tentacle. There are 4 straight radial-
canals which widen as they join the 4 corners of the stomach. The manubrium is wide at its
base and lacks a peduncle. It is about half as long as the bell-height. The 4 oral tentacles
branch dichotomously 8 times, and their tips end in knob-like expansions, armed with bristling
nematocysts. There are thus 256 tips to the oral tentacles in each quadrant.

There are 8 gonads, 2 upon each interradial side of the stomach. These <;onads are dis-
tinctly separated in the 4 principal radii.

The entoderm of the stomach and tentacle-bulbs is orange-yellow or yellowish-green to
brownish-yellow. The gonads are dark-yellow to brown. This medusa is abundant over the
western parts of the tropical Pacific, Fiji, Kllicc Islands. Malay Archipelago West Coast of
Mexico, Acapulco Harbor.

Bougainvillia britannica Forbes.
Plate 17, fig. 8.

lltppvcrtne britannica, FORBKS, 1X41, Annals ami Mag. Nat. Hist., vol. 7, p. 84, plate i. fit;, i.

Bougainvillea britannica, FoRRts, 184X1 British Naked-eveti Medus.T. p. (12, pl.tte 12. fig. I.

Euilfndrium ramosum, VAN BENEDEN, 1844, Mem. Acad. Belgu|ue, tome 17, p. 56, planclir 4; iSdo, F;mn.i I.ittor. Belgique,

p. 112, plates 6, 7.
Bougainvillea ramosa, ALLMAN, 1871, Monog. Tub. Hvdr., p. 311, plate 9, figs. 5-7 (hydroid and medusa). — BOCHU, 1878,

Jena. Zeitsch. fur Naturw., Bd. u, p. 189, taf. 6, fig. 6.
Margtlis ramosa, HAECKEL, 1879, Syst. der Medusen, p. 88.
(?)Ltzusa octocilia (young medusa), HAECKEL, Ibid., p. 80.
Bougainvillia ramosa, WEISMANN, 1883, Entstehung Seiualzellen bei Hydromcdusen, (rn.i, pp. 1 13, 218.— BII.I.ARD, 1906, Bull.

Mus. Hist. Nat. Paris, No. 5, p. 330.

( ?) Bougainvillia ramosa, PICTET, 1893, Revue Suisse Zoo!., tome I, p. II (hvJnnil Ironi Atnhoin.i, M.il.tv Archipellgo).
Lizusa octocilia Dalyell = Bougainvillia ramosa Btihm, HARTI.ACB, 1894, \Visscn. Meercsuntcrsuch. Kciinni. Mern- Kiel,

Helgoland, Neue Folge, Bd. r, p. 190.

\targelis britannica (Forbes), BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, p. 167.
Bougainvillia ramosa, BROWNE, 1898, Journ. Marine Biol. Assoc., N. S., vol. c, p. 187 (hydroid).
Margrlis ramosa, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 138 (citation of papers to 1850).
Margclis britannica, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 754.

Bougainvillia benedenii, BONNEVIE, 1898, Zeit. fur wissen. Zool., Bd. 63, p. 484, taf. 26. fign. 34. ;<; (degenerating hydroid).
(?) Bougainvillia obscura, BONNEVIE, 1898, Bcrgens Museums Aarbog, No. 5, p. 7 (perisarc unringed, 15 tentacles of various

lengths).
Bougainvillia bella, HARTL.AUB, 1897, Wissen. Meeresuntersuch. Kommis. Mcrr<- Kiel, Helgoland, Ncut* Fnli;r. HJ. 2, p. 470,

taf. 15, fig. 7; taf. l6a, figs. 6, 1 1 = B. britannica, Forbes, Hartlaub, 1904, Ibid., B.I. 5, p. 10;.
( .' ?) Bougainvillia muscus, THORNELY, 1900, Zoological Results, A. Willey, Part 4, p. 452 (hydroid from tropical Pacific).

Bell about 12 mm. high and 10 mm. wide. High, dome-like, rounded, with very thick
walls. Each of the 4 radially placed, marginal tentacle-bulbs gives rise to ifi to 24 short
tentacles, about one-fourth as long as the bell-diameter. Each tentacle is provided with a
basal ocellus on its inner (velar) side. The ocelli are black in the adult, but mav In red in
the young medusa.

4 quite wide radial-canals. Manubrium short and wide without a peduncle. The 4
oral tentacles branch dichotomously 6 to 7 times. There are 8 adradial gonads in the ecto-
derm on the sides of the stomach. The ripe eggs are not inclosed in nematocyst capsules

Distinctions l>ct

•,d n.

supcrciltarts.

Marginal tenta-
cles in each of
the 4 radial
clusters.

Character of
stomach.

Character of gonads.

Color.

Length of mar-
ginal tentacles
in terms of
bell-radius (r).

B. superciliaris
L. Agassiz.

10 to 15

Mounted on short,
4-sided, prismat-
ic peduncle.

4 wide interradial. The
eggs develop into plan-

Ulac nil • nit >t

stomach.

Rosin-yellow to
reddish-brown.

1-5 to 2 r

B. britannica
Forbes.

id tci 24

Nn peilunclr.

8 narrow .i,!r.idu! . V
planula- on sides of
stomach.

Golden-yellow.

o.S'

MKDCS.K OF TIIK WORLD.

and do not develop into planulse while still attached to the medusa. The entoderm of the
stomach and tentacle-bulbs is golden-yellow. This species is found oft" the North Atlantic
coasts of France, Great Britain, and Germany. It was taken by Hartlaub at Helgoland,
German Ocean, in June, and by Browne in the Firth of Clyde, Scotland, in June and July.

I found a single mature specimen in the harbor of Eastport,
Maine, on September 19, 1898. It appears, therefore, to be
widely distributed over the North Atlantic. Pictet, 1 893, states
that he identifies the hydroid ol this form from Amboina,
Malay Archipelago.

The hydroid of B. hrittninicn has been commonly called
B. ramosum, and has been well described by Van Beneden and
Allman. The stems are much branched and are 25 to 75 mm.
high; the ultimate branches are for the most part alternate.
The hydrorhiza is branched and root-like, the branches not
anastomosing. The perisarc of the stems exhibits shallow annu-
lations at the origin of the branches. The perisarc extends in
a cup-like form over the sides of the hydranths almost to the
bases of the tentacles, so that the hydranth may be almost
completely withdrawn into the hollow of the cup. There are
about 12 tentacles which are filiform and not very long and
arise in a single zone at the base of the conical hypostome.
The tentacles are carried nearly straight, with every alter-
nate one elevated and depressed. The medusa-buds are borne
singly on moderately long pedicels which arise from the bases
of the hydranths upon the terminal branches of the stems.
There are I to 2 medusa-buds at the base of each hydranth.
When set free the medusa has a bell higher than a hemisphere,
with fairly thick walls. There are 8 marginal tentacles, 2 in
each radial cluster, and 4 short-knobbed, oral tentacles.
Browne, 1898, had a hydroid of this species from the Eddy-
stone, English Channel, which liberated 4,450 medusa; in 3 days.
Wiesman, 1883, finds that the germ-cells of both sexes originate in the ectoderm of the
manubrium of the budding medusa and they do not wander from their place of origin, hut
remain there and become mature in the free-swimming medusa.

Bougainvillia superciliaris L. Agassiz.

Plate 17, fig. I.

Hippocrene bougainvillii, Goi'LD, 1841, Report on Invertebrates of Massachusetts, p. 348.

Hippocrene superciliaris, AGASSIZ, L., 1849, Mem. Amer. Acad., New Series, vol. 4, p. 250, plates 1-3, 53 figs.— STIMPSON, 1853,

Maine Invert. Grand Manan, p. II.
Bougainvillia superciliaris, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 289, 291, figs. 37-39; p. 344, plate 27, figs. 1-7. —

AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 97, figs. 24, 25; 1865, North Amer. Acal., p. 153, figs. 232-240. —

VERRIU., 1873, Report Commiss. Fish and Fisheries U. S., for 1871-72, pp. 328, 733, plate 37, fig. 276.
Hippocrene superciliaris, HAECKEL, 1879, Syst. der Medusen, p. 92.
Bougainvillea superciliaris, WAGNER, 1885, Wirbellosen des Weissen Meeres, p. 73, taf. 2, fign. 5, 8, 9; nan 6, 7.— LEVINSEN,

1893, Vid. Meddel.Nat. For. Kjobenhavn, ser. 5, Bd-4, p. 144.— MAAS, 1893, Ergeb. der Plankton Exped., Bd. 2, K. c.,

p. 69. — HARTLAI B, 1897, Wissen. Meeresuntersuch. auf Helgoland, Heft. I, Abt. 2, p. 466, taf. xvia, fign. I, 5, 7-9, 12;

taf. xvii, fig. 3; taf. xvir, fign. 3, 4, 11-13; ta^- xv' nR- '•
Hippocrene superciliaris, SCHLATER, 1891, Revue des Sci. Naturelles, St. Petersbourg, tome 2, p. 342. — BIRULA, 1896, Annuairc

du Mus. Zool. de 1'Acad. Imperiale des Sci., St. Petersbourg, tome I, No. 4, pp. 330, 339. — GRONBERG, 1898, Zoolog.

Jahrb., Abth. Syst., Bd. ii, p. 462, taf. 27, fign. 5, 6.
Hippocrene superciliaris-^ H. aurea, LINKO, 1904,200!. Anzeiger, Bd. 28, p. 216; 1900, Mem. Acad. Sci. St. Petersbourg,

ser. 8, tome 10, No. 3, p. 6, taf. I, fign. 13-18, taf. z, fign. 19-21 (histology of the ocelli).
Bougainvillia superciliaris, HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 40, I fig.
Bougainvillia superciliaris, NETTING, 1901, Bull. U. S. Fish Commission, vol. 19, pp. 330, 376, fig. 90.
Bougainvillea paradoxica, MF.RESCHKOWSKV, 1879, Annals and Mag. Nat. Hist., ser. 5, vol. 3, p. 177, plate 20.
(?) Hippocrene bougainvillii, BRANDT, 1834, Recueil Actes seances publiques Acad. Imp. Sci. St. Petersbourg, p. 29 (of the

"separate"); 1853, Mem. Acad. Imp. Sci. St. Petersbourg, ser. 6, vol. 4, part 2, p. 393, plate 20= Bougainvillea merunsii,

L. Agassiz, 1862.

Adult medusa. — Bell about 10 mm. in height and about same in diameter. Gelatinous
substance very thick, so that bell-cavity is only about half as deep as the bell-height. There
are 4 clusters of marginal tentacles which are situated at the bases of the 4 radial-canals.

FIG. 86. — Hydroid of Bougainvillia
britannica, after Allman, in Ray
Society, 1871-72.

ANTHOMKDrs.K Hul ( ;\I NVI I.I.I A .

Each cluster of tentacles arises from a large bulbous swelling upon the bell-margin. There
are 10 to 15 tentacles in each cluster. The tentacles are highh contractile and are normally
about two-thirds as long as the bell-diameter. There is a single, very dark-brown ocellus
within the ectoderm ot the inner (velar) side of the base of each tentacle. The velum is well
developed. There are 4 straight, narrow radial-canals, which arise from the 4 radial corners
of the stomach. The stomach is wide and quadratic in cross-section and flask-shaped in
longitudinal outline. It is mounted upon a 4-sided peduncle which is about half as long as
the stomach itselt. There are 4 radially situated, oral tentacles, each of which branches dichot-
omously about 5 times. 1 he mouth is situated at the extremity of a short, simple tube and
the lips are not prominent. The mature genital products are situated within the ectoderm
of the 4 interradial sides of the stomach. There are thus 4 intei radial gonads, above the zone
of the oral tentacles. Ibis species is thus separated from It. li,ll,i by having 4 interradial
instead ot 8 adradial gonads. In the temale, according to Hartlaub, l8i;~, the large eggs
may be seen lying within the ectoderm close to the supporting lamella. As thev gmw larger
they approach the surface, and finally become inclosed in epithelial, nematocyst-bearing
capsules, derived from the ectoderm cells of the stomach-wall. These capsules finally protrude
from the sides ot the manubrium, but each one remains attached to it by a narrow stalk. In
this situation the egg develops into a planula which finally breaks through the walls ot the

capsule and escapes into the water. Hartlaub
showed that although the large eggs were found
in the ectoderm they were derived from the entoderm
of the young medusa. In the young and immature
medusa the supporting lamella does not completely
separate the ectoderm from the entoderm, and the
egg-cells then migrate from the entoderm into the

DO o

ectoderm (see Hartlaub, 180,-, pp. 400, 4-01.* The
sperm originates in the ectoderm and there develops
without migration.

The entoderm of the manubrium and tentacle-
bulbs is rosin-yellow or reddish-brown, and tin-
gelatinous substance of the bell often has a decided
yellowish tinge.

HyJirinl mi,/ \riung medusa. — 'I he hydroid is
usually found attached to rocks, and lives in the
purest sea-water. It grows in clusters, not more than
50 mm. high. The stems give rise to pnmaiy and
secondary branches, which arise somewhat irregu-
larly, though more or less alternately and spirally. The base ot every branch is ringed with from
510 10 annotations. The chitinous sheath which incloses the stem thins out as it passes on to
the polypites and disappears entirely below the circlet of tentacles. Each polypite has from I 5
to 20 long, slender tentacles which arise from a single circlet near the distal end ot the polypite.
Each tentacle is ringed at regular intervals by clusters ot nematocv st-cells. The medusa-
buds arise singly and somewhat irregularly from the sides of the stem, near the proximal
bases of the polypites. Each medusa-bud is inclosed in a thin capsule toimed from the
chitinous ectosarc of the stem. When set tree the young medusa usually has X marginal
tentacles, 2 at the base of each radial tube. The young tentacles are each provided with a
centripetal, ectodermal ocellus. The manubrium is short and tubular and tlu-ie are 4 short,
knob-shaped, unbranched, oral tentacles.

In common with other Arctic medusa? this species appears upon the southern coast of
New England early in April and soon disappears. It is found throughout the summer,
however, north of Cape Cod, and mature individuals are occasionally taken at Eastport,
Maine, in August. This species has been found by Eevinsen, 181;^. off the west coast of
Greenland, and according to Birula, i8g6, it is the most abundant medusa in the White Sea
in July. Hartlaub, 1807, found it at Helgoland, and I.inko, 10.04, found it in Barents Sea,

* It is interesting to observe that other species of the Bougainvillidi, such as ('\i.ri , .I|*M rrt.iin the emhrvos within the ectin-
ilerm of the manubrium until the planula »tji;r. Tins was first observe, I bv Mrr.-si hmvskv, iX?.j, an, I has since been con-
firmed by N. Wagner, 18X5, ami by Hartlaub, 1897.

Fit;. 87. — Bougainvillia su6ffrc///<v7*, after I..

in Mem. Amer, AcaJ. Arts anil Sci
iSatf. Si;le \ii-w uf tnetlusa.

lli-4

MKDl'S.K OF THE WORLD.

where it commonly occurs in the fjords from November until June, but during the summer
it is found about a decree northward of Ekaterinen Harbor, in the cold water. At Helgoland
the medusa is most abundant in February and March.

"Hippocrene inirea" Linko is only an Arctic form of Bougainvillia sitpercilians in
which the oral tentacles branch dichotomously 7 times, instead of 5 to 6 times as is commonly
observed.

I am inclined to believe that Hippocrene boitganivillii Brandt ( = Bougainvillea mertensii
L. Agassiz) from Behrings Sea is identical with B. supercilians.

Fin. 88. — Bougainvillia supcrciliariS) after J>, Agassiz in Mt'in. Amer. Acad.
Arts ami Sci., 1849. Oral view of medusa.

Bougainvillia dinema = B. superciliaris (?).

Bougainvillea litnema, GREENE, 1857, Nat. Hist. Review, vol. 4, p. 246, plate 15, fig. 6.
Margflh zygonema, HAF.IKEL, 1880, Syst. tier Medusen, p. 635.

I believe this medusa to be identical with B. superciliaris. (See tabular synopsis.)

Bell oval with a constricted margin, 10 to 12 mm. high, 6 to 8 mm. wide. The 4 marginal
tentacle-bulbs are kidney-shaped and only half as wide as the intervals between them. Each
bulb bears 10 to 12 ocelli. Only 2 tentacles were observed by Greene and Haeckel on each
bulb, but the others had probably been broken off. The 2 were at the ends of the bulb.

The stomach is roundish, 4-sided, smaller than the tentacle-bulbs, and hardly one-fourth
as long as the bell-height. The 4 oral tentacles branch dichotomously 4 or 5 times. The
gonads are 4 interradial, triangular to egg-shaped swellings on the sides of the stomach.
Stomach, gonads, and tentacle-bulbs yellow.

East coast of Ireland.

Bougainvillia multicilia.

Lizusa multicilia, HAECKF.L, 1879, Syst. der Medusen, p. 81, taf. 6, fig. 13.

Bell three-fourths spherical, with thin, uniform walls, evenly rounded, fi mm. high, 5 mm.
wide. 4 wide, kidney-shaped, marginal tentacle-bulbs, wider than the intervals between
them; each with 10 to 12 tentacles with ocelli, and about as long as the bell-radius. 4 straight.

ANTHOMKIHS.K l« H'CAINVI I.I.I A. I CM

narrow radial-canals. The velum is narrow. The stomach is globular with a constricted base,
but no peduncle. 4 simple, unbranched oral tentacles, each ending in a single nematocyst-
knob. 4 pairs ol gonads on the adradial sides of the stomach. Stomach light-yellow. ( jonads
and tentacle-bulbs orange-yellow. ( kelli dark-red. Straits of Gibraltar in March. This may
be a young form, the shape ot the bell and unbranched condition of the oral tentacles being
indications ot immaturity.

Bougainvillia platygaster.

Hippocrenr plalygasler, HAECKEL, 1879, Syst. tier Mi-ilusen, p. 91.

Bell cubical, flat above. 12 mm. high, 12 mm. wide. The 4 radially placed, marginal
tentacle clusters arise from wide, triangular, basal swellings, which are wider than the inter-
radial spaces between them. Each basal swelling gives rise to 10 to 12 tentacles, which are
shorter than the bell-diameter. The tentacles arise in a single row from their basal bulbs.
Ocelli ( ?) Stomach flat and 4-sided. It is 4 times as wide as high, and without a peduncle.
The 4 short, oral tentacles branch dichotomously 3 or 4 times, giving 8 to 16 short, terminal
ramuli in each quadrant. The 4 wide, rectangular gonads are on the interradial sides of the
stomach. The stomach, gonads, and tentacle-bulbs are reddish-yellow.

Atlantic Ocean, Cape Verde, and Canary Islands.

Distinguished by the wide, triangular, basal bulbs of its marginal tentacles, and its short,
wide, 4-sided stomach.

Bougainvillia xantha Hartlaub.

Bougaim'illia \antlia, HARTLAUR, 1X97, Wissen. Meeresuntersuch. Kommis., Meere Kiel, Helgoland, Neue Folge, Bd. 2, p. 461,
taf. 15, fign. 2, 3; taf. l(ia, fig. 3; taf. l6i, fig. 19.

Young medusa. — Bell rounded, somewhat higher than wide, with thick gelatinous walls.
Height 4 or 5 mm. Bell-margin 4-sided. Each of the 4 radially situated, marginal tentacle-
bulbs gives rise to 9 or 10 tentacles, with small, indifferently developed, brownish-violet ocelli.
4 quite wide radial-canals. Manubnum small, with 4 branching oral tentacles longer than the
manubnum. The oral tentacles branch dichotomously about 4 times and each ends in about
16 nematocyst-knobs. These terminal branches of the oral tentacles are very short in com-
parison with the length of the main shaft of the tentacle. The 4 interradial gonads begin to
develop in specimens about 4 mm. wide. The tentacle-bulbs and stomach are light golden-
yellow to reddish in color. The interradial entodermal lamella is translucent greenish-yellow.
Common at Helgoland from March until the end ot June. No mature specimens have been
seen and the hydroid is unknown. It is closely allied to K. ftiiviJn and may be only a variety
of the latter form. It appears, however, to have more marginal tentacles in corresponding
stages of growth than is seen in R. flnviJn and there are slight color differences.

Bougainvillia carolinensis Allman.

Plate 16, figs. 7 tu 9; plate 17, fig. 7.

Hifiporrfne tarolinfnsn, McCfiADV, 1857, Gynin. Charleston Harbor, p. hi, plate 10, figs. 8-10.

Margelis carolinrnsis, AI.ASSIZ, I,., 1862, Cont. Nat. Hist. l; . S., vol. 4, p. 544. Anssi/, A., 1865, North Amir. Acal., p. 156,

tigs. 241-248.— HAECKEL, 1879, Syst. iler Meilusen, p. 89.
Bou^ainl'illia ctirolinensis, AI.LMAN, 1871, Monog. Tubul. Hvdroids, p. 316.

Hougiiimi/lia carolinensis, NUTTINC., 1901, Bull. U. S. Fish Commission (or 1X99, vol. 19, pp. 350, 376; figs. 5, 89.- HARCITT,
1904, Bull. I'. S. Bureau of Fisheries, vol. 24, p. 39, plate 2, fig. 4; 1901, American Naturalist, vol. ^, p. ^83, fig. 46.

AJult nit'Jnsii (plate 17, fig. "). — Bell dome-shaped and 4 mm. in height. Bell-walls,
especially at apex, very thick. There are 4 radially arranged clusters of marginal tentacles
that arise from 4 small, bulbous, basal swellings. 7 to g tentacles in each cluster. The longest
of these tentacles attain a length about equal to the bell-height. The tentacles are slender
and quite stiff, and their tips are usually curled downwards in a semicircle. A large ectoder-
mal ocellus is situated at the base of each tentacle upon the ventral (centripetal) side. The
velum is well developed. There are 4 straight, narrow, radial tubes. The manubrium is
long and narrow, and extends for about half the distance from the inner apex ot the bell-
cavity to the velar opening. Near its distal end it is somewhat wider than at its base. The
mouth part, however, is a simple, narrow tube. 4 oral tentacles arise from the 4 radial sides

MKDrs.E OF THE WORLD.

of the mouth-part of the manubrium. Each one of these branches twice dichotomously, thus
giving rise to 4 tentacle tips in each quadrant. The gonads are found in the ectoderm of the
manubrium, where they occupy longitudinal, interradially situated, swollen regions.

In the Tortugas and Charleston Harbor examples of this species, the entoderm of the
tentacle-bulbs and of the manubrium is of a delicate sage-green. The gonads are cream-
colored, and the tentacular ocelli dark-brown or black. In northern examples of this medusa,
found at Newport, Rhode Island, and Naushon, Massachusetts, according to A. Agassiz, the
tentacle-bulbs are colored with brilliant red, surrounded by a green edge bordered with bright-
yellow, and the digestive cavity is brick-red or green. No such brilliant coloration has been
observed by me in southern examples of this species.

The hydroid stock has been figured by A. Agassiz, 1865, p. 158. It occurs very commonly
upon Fuciis vesifulosus, where it grows in dense clusters. The main stems are often 230 to
270 mm. in length. They taper gradually from base to summit and branch profusely and pin-
nately. The side branches do not often give rise to secondary branches. Great numbers ot
ringed pedicels, which terminate in polyp-heads, arise from the sides of the main stem and also
from the branches. The polypites are fusiform and possess a single circlet of about 15 long,
slender tentacles near the distal extremity. The mouth is situated at the apex of a slight, conical
proboscis. Medusa-buds make their appearance anywhere all over the stems, where they
appear either singly or in clusters. When set tree the young medusa is about 1 .7 mm. in height.
The bell is pyriform in outline, and the gelatinous substance quite thick. There are 8 tentacles,
2 at the base of each of the 4 radial tubes. The tentacles are all provided with basal ocelli.
The manubrium is small, slender, and tubular, and there are 4 short, unhranched, oral ten-
tacles.

The general color of the stems of the hydroid is of a horny green and the polypites are
often of a delicate rosy tint.

This medusa is very abundant throughout the summer in Charleston Harbor, South Car-
olina. It is much rarer at the Tortugas, Florida. The brilliantly colored, northern variety is
said to be quite common upon the southern coast of New England, but it has never been
found north of Cape Cod, Massachusetts.

Hargitt, 1901 (Biol. Bulletin, Woods Hole, vol. 2, p. 228), discovered a remarkable twin
sport, apparently of this medusa. It had 2 manubria fused at a common base so as to form a
common stomach. Each "twin " had a complete bell-margin, only the sides ot their bells being
fused one to the other.

Bougainvillia niobe Mayer.
Plate 18, figs. 1-3.

Bougainvillia niohe, MAVF.R, 1894, Bull. Mus. Comp. Zool. at Harvard College, vol. 25, p. 236, plate I , fig. 2: Ifiiil., vol. 37, p. 42 ;
1904, Mern. Nat. Sd. Brooklyn Institute Museum, vol. i, p. 11, plate z, figs. 14-ijr.

Adult medusa (plate i8,fig. I ). — Bell 6.75 mm. high and 4.8 mm. in diameter, with vertical
sides and thick, gelatinous walls. 4 radially situated clusters ot marginal tentacles, each com-
posed of about 8 tentacles about as long as the bell-height. On the velar side of each tentacle,
near the bulbous base, there is a dark-colored ocellus. 4 straight, narrow radial-canals.
Manubrium wide, flask-shaped, and cruciform in cross-section. About half as long as the
height ot the bell-cavity. 4 radially situated clusters of oral tentacles, each of which branches
dichotomously 4 times. These are very flexible and their distal ends are knobbed.

Medusa-buds arise from the 8 adradii of the manubrium, the youngest buds being im-
mediately under and on both sides of the point of entrance of each radial-canal, while older
ones are found farther down the sides of the stomach. The young buds are small, ovoid
vesicles contained entirely in the ectoderm. The limiting membrane, between the entoderm
and ectoderm of the manubrium in the neighborhood of these buds, is entire (plate 18, fig. 3),
and I can find no evidence of cells passing through it. Indeed, throughout the future develop-
ment of the buds, the entoderm remains inert and its limiting membrane unbroken. The
wall of the vesicle consists of 2 layers of cells: an outer epithelium of ectodermal cells, des-
tined to give rise to the ectoderm of the hud, and an inner layer of somewhat larger cells
destined to give rise to the entoderm of the budding medusa (plate 18, fig. 2). These latter
cells are entirely incased by the ectoderm of the parent. They may, however, have been

PLATE 18.

Fig. I. Bougainvillia niobc, Nassau Harbor, Bahamas, March 18,

Fig. 2. Roiiaiilnvillla niobc, section ot a young medusa-bud upon the ecto-
derm of the manubrium. r, central mass of degenerating cells;
c <• t, ectoderm of the manubrium of the parent medusa; c n t,
(brown) entoderm of the parent medusa; e c t b, (gray) ectoderm
of the budding medusa; e n t b, (green) entoderm of the budding
medusa; s I, limiting membrane between the ectoderm and ento-
derm of the manubrium ot the parent medusa.

Fig. 5. Bougainvillia niobc. Cross-section of the medusa showing the
budding region of the manubrium. Successive stages in develop-
ment of buds are shown in hi b4. Colors as in figure 2.

Fig. 4. R/it/iki'/i flea/ins, mature male. Tortugas, Florida, July I, 1899.

Fig. 5. Rathkea octof>unct<it(i var. grata. Newport Harbor, Rhode Island,
April 24, 1897.

Drawn by the author from living and preserved specimens.

PLATE 1 8

' Ct.

ent.

"•/. l,.

si.

i nt . l,.

2

.\\THuMi:i>[ S.K HnrCIATXVII.i.IA. 1C.7

derived at an earlier stage from the entoderm of the parent medusa and have migrated into
the ectoderm. Careful search has, however, failed to reveal evidence of any such migration,
and in default of evidence to the contrary, we assume that these cells niiiv he ectoilermal in
origin, although destined to produce the entoderm of the hud.

The central cavity of the vesicle is partially rilled with a loose mass of nuclei and cell ma-
terial showing little or no trace of cell boundaries (r, plate 18, fig. 2). These are present at all
stages of the developing hud, partially filling the gastrovascular cavity, hut they appear to
degenerate in later stages, and it seems possible they serve to nourish the bud in its urowth.
This supposition appears the more probable from the fact that the gastrovascular ca\itv of the
bud is never in connection with that of the parent medusa.

\Ve have, therefore, a graded series of phenomena in the asexual production of medusa-
buds by hyd mined usx. Where the ectoderm and entoderm are both thin-layered and of about
equal thickness, such as in Stirs in and F.uclifilr,t,i p,nnj<,\iia, both entoderm and ectoderm
take an equal share in the formation of the bud. In forms such as Ruthk,,: 'i.t't[>un<-tntti and

•* ' ° '•

1 i I

1 . i m

S:

-V Ao,,"

Sq. 89. 9°-

l-'n,. Sij. Hvilrohl of Hvi<i!<i:niillni carolinfmis, after A. Agassi/, in North American Acalepluf,

showing an eiilar^eil view of .1 part of >tem liearin^ medusa-buds.
I']'.. 90. — Boutylim'illiil niuhf, from life, liy the author. Tortuy.is. I;lon.la, June 2", 1908.

Lizzin fliiftrirt-tli-i, according to Chun, Bibliotheca Zoologica, 1895, a different condition is
observed, for the medusa-buds are formed entirely within the ectoderm of the parent, although
the gastiovascular cavity of the bud finally acquires a connection with that of the parent, tin
entoderm of the bud becoming continuous with that of the parent manubrium.

In Bougainvillia nin/ii-. however, the ectoderm is very thick and the budding medusae
are developed within it alone, the entoderm of the parent medusa remaining inert and passive
during the growth of the bud, and no connection ever being established between the gastro-
vascular cavities of the bud and the parent. This result may he regarded as due to a gradual
process favored by the thickness of the ectoderm, which prevented the deep-lying entoderm
from taking an equal share in the formation of the bud. until fiuallv it remains passixi-
throughout the period of the formation of the bud. Medusa? produced from ectoderm alone
may, therefore, be phylogenetically homologous with medusae produced by the more primi-
tive and universal cooperation of both ectoderm and entoderm.

The entoderm of the manubrium and tentacle-bulbs of K. niobe is rosin-colored, all other
parts being transparent. There is a dark-colored ocellus on the velar side of each tentacle
near the basal bulb. This medusa is found in Nassau Harbor, Bahamas, in April.

168

MF.TH'S.-E OK THE AVORLD.

Bougainvillia pyramidata.

Hipforrrur f\ramiilaia, FORBES AND GOODSIR, 1853, Trans. Roy. Soc. Edinburgh, vol. 20, p. 312, plate 10, fie. 4. — HAFCKH ,
iSSo, Syst. .ICT Mcilusrn, p. 635.— BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 75?.

Bell semiglobular, 2.5 to 6 mm. wide and equally high. Bell-walls thick and gelatinous
substance at apex well developed. The 4 marginal tentacle-bulbs are very small and globular,
and each one bears 6 to 8 tentacles which have ocelli and are somewhat shorter than the
bell-diameter. The stomach is mounted upon a short, conical peduncle. It is cruciform and
its 4 lobes extend along the entire length of the peduncle. The mouth is a circular opening.
The 4 oral tentacles branch twice dichotomously. The gonads are upon the basal margin
of the stomach and its lobes. The radial-canals are straight and narrow. Stomach, gonads,
and tentacle-bulbs orange-yellow to reddish-brown. Tentacular ocelli black. Found off
the west coast of Scotland and off the south coast of Ireland.

Bougainvillia nordgaardii.
Margtlis nordgaarJii, BROWNF, 1905, Bergens Museums Aarbog, No. 4, p. 14, plate 2, fig. I, plate 3, figs. <, 6.

Bell 4 mm. wide, 4 mm. high. Bell-walls uniform and of moderate thickness. Bell
dome-like, evenly rounded. 5 to J tentacles in each of the 4 radial bulbs. No ocelli. Ten-
tacles somewhat longer than the bell-diameter.
Velum narrow. Stomach half as long as the depth
of the bell-cavity. 4 oral tentacles, 4 or 5 times
dichotomously branched. 4 wide, mterradial,
gonads on the sides of the stomach. Gonads and
tentacle-bulbs yellowish-brown (in formalin).
Byfjord, near Bergen, Norway. Distinguished
by its lack of ocelli.

For details see tabular synopsis of medusae of
Bougainvillia.

Bougainvillia flavida Hartlaub.

Bougainvillia

imillin faritla, HARTLAUB, 1897, Wissen. Meeresuntersuch.
Koinmis. Deutsch. Meere Kiel, Helgoland, Neue Folge, Bil.
i, p. 456, taf. 14, figs, i-io; taf. 15, figs. 4-6, 8.

The mature medusa is slightly higher than a
hemisphere with an evenly-rounded bell. The
gelatinous substance is thick, so that the bell-
cavity is only a little more than half as deep as
the bell-height. The bell is 2 or 3 mm. wide. There
are 4 radially placed clusters of marginal tentacles
with not more than 6 tentacles in each cluster.
These tentacles are about as long as the bell-
diameter and each one has a large ocellus upon its
velar side near the base of the cluster. The 4 com-
mon, basal bulbs are swollen, semiglobular, and
well developed. 4 quite wide, straight radial-canals
and a narrow, simple, circular vessel. Manubrium small, conical, half as long as the depth of
the bell-cavity. Without a peduncle. The 4 oral tentacles branch dichotomously about 3
times. With mterradial, undivided gonads. Entoderm of manubrium and tentacle-bases
orange-yellow.

The mature medusae are found at Helgoland, German Ocean, in August and September.
Hydroid. — The hydrorhiza is net-like, creeping, and yellowish-brown in color. The
stems arise at quite infrequent intervals and are unbranched, very slender below and swollen
above at the polypite. They are 5 to 7 mm. high and light gray-brown in color. The hydranths
are elongate, club-shaped, with 8 to 10 tentacles in a single zone around the mouth. The
medusa-buds are produced upon branching processes which arise from the sides of the stem
near the base of the polypite, or from points lower down upon the side of the polypite

!i',.<ji. Bougainvillia nordgaardii, after Browne
Hergens Museums Aarbog, 1903.

\.VI Iln.Ml.lH s.K liiil (IA1NV1U.IA. Iti'.l

itself. There may be as many as 6 medusa-buds upon one of these branches, and there are
usually several branches producing medusae. The buds are oval at first, but later they
become spherical.

When set free the medusa's bell is only 0.3 to O.6 mm. wide, li is irregularly glob-
ular with very thick walls and an irregularly shaped bell-cavity. Tbeie are 4 penadi.il basal
bulbs, each of which gives rise to a single, marginal tentacle. These are long and lack ocelli.
4 simple, unbranched, oral tentacles, and 4 wide radial-canals, with an axial canal above the
stomach. The exumbrella is besprinkled with nematocysts as is usual in young medusx.
The tentacle-bulbs and manubrium are reddish-brown by reflected light.

The hydroid is found at Helgoland attached to living or dead shells during the summer.
Both hydroid and medusa are described in detail by Hartlaub, 1897.

Bougainvillia autumnahs Hartlaub.

I'laU- id, figs. 4 and 5; plate 17, figs. 3 and 4.

KI ROPKAN MKDI SA.

Bottgainvillia autumnalist HARTLAI B, iS^y, \\issen. Merrrsuntersuch. Knmmis. Mrrrr Kn-1, Hrlgnland, NVur Fnl^r, Bd. 2,

p. 461;, taf. l <;, fign. 11-13.
Mti'gt'lii auturririuli), BROWNE, !9Od, Trans. Linnean Soc. London, ser. 2, Zoo]., vol. 10, p. 170.

AMKRKAN MKDUSA.

Bougainvillia gikbsi, MAYKR, 1900, Bull. Mus. Comp. Zool. at Harvard Collrg'-.. ?ol. ;-. p. ;. pl.ii>' 4. hi; 14, t v HAM. ITT,
1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 40; 1901, American Naturalist, vol. 35, p. 583.

Bell rounded, 2.5 mm. wide, somewhat higher than a hemisphere, with thick, gelatinous
walls. 4 bulbous, rounded, marginal tentacle-bulbs, each of which gi\es rise to 3 to <i ten-
tacles. 4 narrow, straight radial-canals. Manubnum short, wide, 4-sided, urn-shaped;
not half as long as the depth of the bell-cavity. 4 oral tentacles which turn upwards and
branch dichotomously 2 to 5 times. The ends of the oral tentacles are simple, without terminal
knobs. 4 interradial gonads in the ectoderm of the stomach-wall. Tentacle-bulbs yellowish
to greenish-yellow. Manubrium green or brown. Hydroid unknown.

Found by Hartlaub at Helgoland, German Ocean, from the end of August to the end
of November. Browne found it off the west coast of Ireland and in the English Channel.

I consider this medusa to be identical with B. %ibbsi Mayer, of the southern coast of
New England. It is possibly separated by uncertain and variable color differences. A
description of the American form is herewith presented:

Adult iiirJnm (plate 16, fig. 5; plate 17, fig. 4). — Bell about 4111111. in height and 3.8 mm.
in diameter. Gelatinous substance very thick, so that the bell-cavity is only about hall as
deep as the height of the animal. There are 4 clusters of marginal tentacles which arise from
4 large, bulbous swellings situated at the bases of the 4 radial-canals. Each bulbous swelling
gives rise to 4 or 5 slender tentacles. There is a single, dark-brown ocellus at the base of each
tentacle upon the centripetal (velar) side. The velum is small. There are 4 straight, narrow
radial-canals. The manuhrium is wide and cruciform in cross-section, and the radial-canals
arise from the 4 coiners of the cross. The manubnum is short and docs not extend quite
half the distance from the inner apex of the bell-cavity to the velar opening. The mouth is
situated at the extremity of a short, tubular neck and there are no prominent lips. 4 radially
situated, oral tentacles arise from the sides of the neck of the manubrium. Each of these
branches dichotomously about twice. The gonads are developed upon the 4 interradial
sides of the stomach, and in the female the ova are large and prominent. The manubrium
is pearl-colored or of a delicate green. The entodermal cores of the tentacle-bulbs are red,
surrounded by a delicate yellow -green. The supporting lamella of the bell often displays a
faint greenish tinge.

I'nniii; mi-iliisa. — In the young medusa there are hut 8 tentacle.s, 2 from each tentacle-
bulb. The bell is a little higher than a hemisphere and the gelatinous substance is not very
thick, being of about uniform thickness everywhere, instead of being very thick at tin aboial
pole as in the adult. The manubrium is short and quadratic and there are 4 short, un-
branched, knob-shaped, oral tentacles. When the medusa is about $ mm. in height, the bell
is still hemispherical. The manubrium is wide, shallow, and quadratic, and the oral tentacles
branch once dichotomously. About } to 5 marginal tentacles arise from each tentacle-bulb.

170

MKIHS.K <IF THK WORLD.

This medusa is found in Newport Harbor, Rhode Island, United States, from July until
( )ctober. I consider it to be identical with Haitlaub's B. autumnalis from Helgoland, German
Ocean.

This species is distinguished from Kougainvtllia carolinensis by the greater height and
lesser width of its bell. Also in B. carolinensis the manubrium is long and slender, while in
B. tiutuiiiiiiilis it is short and wide. The manubrium of B. curolinensn is widest at about the
middle of its length, while that of B. antuiiinnlis is widest at its proximal base.

Bougainvillia prolifera.

l,i a-, i prolifera, VON LKNDKNFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 589, plate 23, figs. 38, 39.

Bell evenly rounded, j mm. high, 2.5 mm. wide. 4 radial clusters of marginal tentacles
which arise from large, "scrota-shaped" basal bulbs; 5 tentacles upon each bulb, 20 in all.
These tentacles are about half as long as the height of the bell. 4 simple radial-canals.
Stomach small, nearly cubic, with 4 unbranched, simple, oral tentacles about two-thirds as
long as the length of the stomach. These end each in knob-like clusters of nematocysts.
Medusa-buds develop at the base of the manubrium and these give rise also to medusae which
appear as buds at the time when the originally budding medusae are set free. Tentacle-
bulbs and stomach brown, other parts colorless.

Port Jackson, New South Wales; March to April.

This medusa exhibits every indication of immaturity, and it is probable that the oral
tentacles finally become branched.

Bougainvillia maniculata Haeckel.

Bougainvillea manirulala, HAECKKL, 1864, Jena. Zeitsch. fur Naturw., Bd. i, p. 340; 1879, Syst. der Medusen, p. 89, taf. 5,
fign. 4, 5.

Bell almost spherical, 1.5 mm. wide, 1.5 mm. high with uniform, fairly thin walls. Velum
very wide. The 4 marginal tentacle clusters are hand-shaped ; each with 4 very short,

92.

93-

Fu.. 92. — Brjii^iinrilliii maniculata, after Hacckcl, 1879.

FK.. 93. — Bongainvillia rugosa, after S. F. Clarke, in Mem. Boston Soc. Nat. Hist.

finger-like tentacles, with an ocellus on the inner side of each. The stomach is spherical,
with a narrow base, and a long, conical throat-tube which projects beyond the velar opening.
There are 4 long, slender, oral tentacles, which branch twice dichotomously at their outer
ends, thus ending in 4 very short, knobbed tips. These oral tentacles are each about as long
as the manubrium itself.

There are 4 interradial gonads on the sides of the stomach. The stomach, gonads, and
tentacle-bulbs are violet-gray. Found by Haeckel in the Mediterranean at Villafranca and
Nizza.

ANTHOMKDIS.E — Uol (iAINVI I.I.IA . 171

The medusa is distinguished by its thin-walled, almost spherical hell, its wide velum.
and its very long oral tentacles which branch only at their outer ends. The Mask-shaped
stomach and small size are also characteristic. On the whole it presents the appearance of
a degenerate torm.

Bougainvillia rugosa Clarke.

Plato 17, fig. 2..
Bougainvillia rugosa, CLARKI., |88>, Mem. Huston Soc. Nat. Hist., vol. 3, p. 140, plate 8, figs. 11-14.

) tniti^ ini-1/iisn (plate 17, tig. 2). — Umbrella somewhat pynlorm. 12 marginal tentacles, $
at the base ot each radial tube. Tentacles all of equal size. According to Clarke, ocelli air
developed only at the bases ot those 2 tentacles which are first and second as one passes
around the bell from left to right, but I found a specimen with 12 ocelli, one at the base nf
each tentacle. The velum is well developed. There are 4 straight, narrow, ladial tubes.
The manubrium is short and thick, and there are 4 short, unbranched, oial tentacles.

H \drnid. — The hydroid stock attains a height of 75 mm. There are numerous irregu-
larly arranged branches. Some of the branches which arise near the proximal end arc about as
long as the main stem, but most of them are short and delicate. The branches bear secondary
branches which give rise to 3 or 4 terminal ramuli. The hydroid is rooted by a creeping stolon.
The polypites are fusiform and are protected by an expansion ot the pensarc which bears a
number of annulations. There are 8 to 10 short tentacles. The medusa-buds are borne upon
the ultimate branches immediately below the proximal bases ot the polypites. I lie hydroid is
light-brown.

It was found by Clarke, 1882, growing upon .//< \rnildluin at Hampton Roads, Virginia,
and in the southern parts of Chesapeake Bay. A medusa believed to be of this species was
found by me in Charleston Harbor, South Carolina, on December 17, 1904.

Bougainvillia trinema.

Margin trinemti, VON LKNDKNFELD, 1884, Proc. Linncan Soc. New South Wales, vol. 9, p. 918, platr 41, fig. i }.

An immature medusa from Sydney Harbor, New South Wales, Australia. The bell is
oval, rounded, 2 to 3 mm. wide. There are only 3 tentacles in each of the 4 clusters of marginal
tentacles. These are about half as long as the bell-height. Each of the 4 oral tentacles ends
in 3 ( ?) small, equal branches about one-third as long as the shaft of the oral tentacle. Ento-
derm ot stomach and tentacle-bulbs dark yellow. This may possibly be the young of II. fulva
Agassiz and Mayer, 1899, but von Lendenfeld's description and figure are too imperfect to
determine this point.

Bougainvillia frondosa Mayer.

Plate 16, fig. 6.
Bougainvillia fronJosa, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard Collide, vol. 37, p. 41, fig. 5, plati- -,.

The bell is higher than a hemisphere with relatively straight, vertical sides, and about
2 mm. high. There are 4 radially placed clusters of marginal tentacles, which are situated
upon bulbs at the ends of the 4 radial-canals. Each bulb gives rise to but 2 tentacles, so that
there are only 8 in all. There are no ocelli at the bases of the tentacles. The velum is narrow.
There are 4 straight, narrow radial-canals and a simple, narrow ring-canal. The manubrium
is cruciform in cross-section, short, thick, and flask-shaped. It extends about half the dis-
tance from the inner apex of the bell-cavity to the velar opening. There are 4 radially placed.
oral tentacles, each of which branches dichotomously 2 or 3 times. Planulx develop upon
the sides of the stomach near, and on both sides of, the 4 principal jadn. I at first consid-
ered these to be parasitic Zooxanthellae, but further studies have shown that they are pl.innl.i
Hartlaub, 1897, observes a similar phenomenon in Bougainvillia superciliarii, etc., and
Maas, 1904, cites it in (*.\t<m mgritinn (Result. Camp. Prince de Monaco, fasc. 28, p. 8. plate
i, tigs. 3, 4).

The entoderm of the manubrium and tentacle-bulbs is pinkish-white and the tentacle tips
are turquoise. This medusa is occasionally found at Tortugas, Florida, in June to Julv.

172 MKDUS.E OF THE WOULD.

Bougainvillia obscura Bonnevie.

li'iugaim-illia ohscura, BDNMVIK, 1899, Bergens Mus. Aarbog for 1898, No. 5, p. 7, taf. i, fig. 4, 40.

The hydroid is small, about 20 mm. high. The hydrocaulus of the trophosome is about
4 mm. high, with almost opaque perisarc without foldings or rings. The polypites are about
2 mm. long, nearly cylindrical and with about 15 very long tentacles. The medusae are set free,
and are produced either singly or in small clusters upon branched processes immediately under
the hydranths.

Found at Christiania. Color ( ?) Hydrorhiza ( ?) Description of medusae ( ?) The
species is said to differ from other Bougainvilliae in its smooth, opaque perisarc, and in its long
tentacles. It will be difficult to rediscover this form from its very meager description.

Bougainvillia glorietta Torrey.
Bougainvillia ghriftta, TORREY, 1904, Univ. California Publications, Zool., vol. 2, No. i, p. 7, figs. 2, 3.

Stems branched, rising trom a creeping hydrorhiza in clusters to the height of 20 to 30 mm.
The stems and branches often twine about each other. Perisarc smooth, or wavy, covered with
particles of detritus. Terminal polypites largest, with 20 to 25 tentacles in 2 or 3 irregular
whorls, the outermost being the shortest. Tentacles highly contractile. Gonophores in groups
of 2 or 3 on branches or hydranth stalks, mounted on short pedicels. The well-developed
medusae within the gonophores have 4 pairs of marginal, 4 simple, oral tentacles, and 8 ocelli.
Color ( ?) Found at San Diego, California.

Distinguished by its smooth, unannulated perisarc and twining stems. Free medusa is
unknown.

Genus NEMOPSIS L. Agassiz, 1849.

A>wo/m.<, AGASSIZ, L., 1X49, Mem. Anier. Acad., New Series, vol. 4, p. 289; 1862, Cont. Nat. Hist. U. S., vol. 4, p. 34^. —
McC-RADY, 1857, Gyrnn. Charleston Harbor, p. 58. — AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 98; 1865,
North Amer. Acal., p. 149. — HAF.CKEL, i879,Syst.der Medusen, p. 92.— BROOKS, 1883, Studies Biol. Lab. Johns Hopkins
University, vol. 2, p. 468. — HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 37.

This genus was founded by L. Agassiz, 1849, t°r Nemopsis bachci of the Atlantic coast of
the United States south of Cape Cod.

GENERIC CHARACTERS.

Margelmae with 4 clusters of marginal tentacles, one at the extremity of each of the radial-
canals. There are two sorts of tentacles in each cluster, a median pair of club-shaped tentacles
with swollen extremities studded with nematocysts; and on both sides of this median pair
there are a number of simple, filiform tentacles. The 4 radial lobes of the stomach extend out-
ward along the radial-canals and the gonads are developed along the sides of these lobes. The
hydroid is a Bougainvillia. In the Nemopsis hydroid, however, the medusae arise from the
sides of the naked polypites, not trom the branches of the stems of the hydroid, as is usual in
other species of Bougainvillia.

In common with Bougainvillia the medusa of Nemopsis has 4 dichotomously branching,
oral tentacles. Its most striking feature is the wide extension of the 4 radial pouches of the
stomach along the 4 radial-canals. This is, however, not of generic value, for it occurs to a
greater or less degree in Bougainvillia. The sole distinctive generic feature of N emopsis is the
presence of two distinct sorts of tentacles upon each marginal bulb. L. Agassiz, in his first
brief notice of this genus in 1849, called attention to its peculiar club-like median pair of ten-
tacles and presents a good figure of them, but his description is not wholly correct, for the ocelli
are not upon the ends of the tentacles, but are on the axial (inner) side of each tentacle adjacent
to the pad-like base.

Haeckel, 1879, considers Nemopsis to be identical with Favonia of Peron and Lesueur,
1809; nevertheless he adopts the name Nemopsis, although maintaining the precedence of
Fnvnnin. I agree with VanhSfFen, 1891, and consider that Favonia is certainly not equivalent
to Nemopsis. Blainville's (1834, Man. d'Actinologie) copy of Peron and Lesueur's figure of

ANTHOMEDU&E— NEMOrSIS. 17.'!

Favonia and his briet and unsatisfactory description are only confusing. It is evident that the
figure is very erroneous, and it appears to hear a closer relationship to I.\'»ni<i>,-<i than tn
Nfinrjpsis. No marginal tentacles are shown, and the flat, hell-shaped disk is wholly unlike
any known species of Nemopsis.

favonin octonfma Peron et Lesueur is probably an imperfect specimen of some species
of Lymnorea. Haeckel calls this "Nemopsis favrjnin."

I can distinguish no difference between HaeckiTs A', -WO/MM " li,-t,-i nnnnn " and A'.
/>iii-lii-i ot our coast. It was described from an isolated specimen found on the coast of \<>i-
way, and falls within the common limits of variability of N. bachci in all respects. It is
possibly an individual which may have drifted across the Atlantic in the trend of the winds
and currents.

The genus Nftnnpsis is evidently derived from the more generali/.ed l}nn!'<iiii;-illi-i in
which the marginal tentacles have become speciali/.ed into two sorts and the 4 radial corners
of the stomach have extended far outward along the radial-canals. The hydroid remains
identical with that of Bougainvillia.

Nemopsis is one of those exceptional Anthomedusae in which the gonads have migiati-d
outward from the sides of the stomach along the radial-canals. J)i>Min,'in,i among the Tiariiue
is another instance ot the same sort. In the medusa; we frequently find that the hard and fast
distinctions which Haeckel attempts to apply to the separation of families do not hold in
nature. Thus the gonads are often more or less interradial in the Codonidae, or more or less
upon the sides of the manubrium in the Thaumantiada;; the lithocyst-bearing medusa l:ntn>i,i
arises from a hydroid having the superficial appearance of one of the Tubularian order, and
Rntlikca fascicultita is the only medusa of the Oceanidae with 4 frrr/iJinl gonads.

Nemopsis bachei L. Agassiz.

Plate 17, figs. 5 anil 6.

Ncmopsn bachei, AGASSIZ, L., 1849, Mem. Amer. Acad., New Scries, vol. 4, p. 289, i fig. 1862, Cont. Nat. Hist. I . s.,
vol. 4, p. 345. — AGASSIZ, A., 1862, Proc. Boston -Soc. Nat. Hist., vol. 9, p. 98, figs. 26, 27. — 1865, North Amir. A. ll.,
p. 14*), figs. 227-231 . — VKRRILL, 1873, Report Commis. Fish and Fisheries U.S. for 1871-72, pp. 454, 733- — HAF< KM,
1879, Syst. der Medusen, p. 93. — BROOKS, 1883, Studies Johns Hopkins Univ. Biol. Lab., vol. 2, p. 468 (hvdrmd and
young medusa). — BROOKS, 1890, Handbook of Invert. Zoo]., p. 40, fig. 21. — NI'TTING, 1901, Bull.U. S. Fish Cnmmis-
sion for 1899, vol. 19, p. 375, fig. 88. — HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 41, I fig.; 1901 , Ameri-
can Naturalist, vol. 35, p. 583, fig. 47; 1901, Biol. Bulletin, Woods Hole, vol. 2, p. 227.

Vfinnftsi! gibbuii, McCfiADV, 1857, Gymn. Charleston Harbor, p. 58, plate 10, figs. 1-3, non figs. 4-7.

Favonia bachei, HAECKF.L, 1877, Prodrom. Syst. Medusen, No. 103 (unpublished).

.Vemopsis heirronema, HAF.CKEL, 1879, Syst. der Medusen, p. 93, taf. 5, fign. 6-9 (isolated specimen from coast of Norway).

Adult medusa (plate 17, fig. 6). — Bell about II mm. high, being higher than a hemi-
sphere, with relatively straight sides and flat top. There are 4 radially situated clusters ot
marginal tentacles. Each cluster arises from a cleft, pad-like base and contains about 14
tentacles. The median pair of tentacles are short, slender, and terminate each in a knob-
like cluster of nematocysts (plate 17, fig. 5). These median tentacles are flanked by a long,
highly contractile pair which contain reddish-brown pigment in their ectoderm. On either
side of these brown tentacles there are 5 transparent, highly contractile, filiform tentacles.
When swimming the medusa often carries its tentacles with their shafts extending straight
outward at an acute angle with the sides of the bell, while the extremity ot each tentacle bends
sharply downward at a right angle, recalling the appearance presented by the tentacles ot
Gonionemus. There is a dark-colored ectodermal ocellus at the base of each tentacle on the
lower (velar) side. The bell-cavity is about three-quarters as deep as the bell-height. There
are 4 straight, narrow radial-canals and a slender circular vessel. The velum is well developed.
The gonads are flat and ribbon-like, with curtain-like folds, and they extend down the radial-
canals, along the sides of the 4 stomach-pouches, which project about two-thirds the distance
from the sides of the manubrium to the tentacle-bulbs. The manubrium is short, with 4
simple, quadratic lips, and with 4 radially situated clusters of oral tentacles. Each of these
oral clusters arises from a single shaft which bifurcates from 5 to 7 times, terminating in 32
to 128 small, nematocyst-covered knobs. The entodermal cores ot the oral tentacles consist
of chordate cells. Gonads, stomach, and tentacle-bulbs dull milky-yellow to orange. This

'4

174 .M 101)1 S.K OK THK WORLD.

medusa is very abundant in autumn and winter along our Atlantic coast near the mouths
of large hays into which pure ocean water has free access. It is not usually found in brackish
water far from the ocean. It is found at Woods Hole, Massachusetts, and is common in Long
Island Sound as far west as New Haven; in the mouth of the Chesapeake; in Pamlico Sound,
especially near the inlets, and in the mouths of all southern harbors, in winter, as far south
as Fernandina, Florida. It is also found off the coast in the ocean. It has not been seen
north of Cape Cod, nor upon the southern Florida coast. Very often the medusa is seen
with all of its marginal tentacles broken off short, leaving only rounded stumps, this condition
being commonly found in animals living in brackish water or far from the ocean. N. " hetero-
iifnui" HaeckeJ, 1879, appears to be identical with our American TV. b/ichei.

HyJronl <inJ young medusa. — McCrady, 1857, is mistaken in his identification of the
hydroid of this medusa, for his hydroid is Margelopsis, The hydroid stock was found by
Brooks, 1883, in Newport River, North Carolina, growing upon a piece of submerged wood.
It is a Bougainvillia. The stems oi the hydroid are about 25 mm. in height. Each mam
stem gives rise to 3 or 4 short alternating branches, and these as well as the main stem end in
hydranths, which are sharply separated from the stem by means of a fold or collar. The
stems are covered by a thin, transparent, horny ectosarc which extends almost, but not quite,
up to the basal collar of the polypites. There are 2 or 3 irregular annulations upon the ectosarc
of the side branches, close to the mam stem. Each polypite bears 24 long, slender tentacles,
which arise in a single circlet near the distal end of the hydranths. The mouth is situated
at the extremity of a well-developed proboscis which is tunnel-shaped and sharply distin-
guished from the body of the hydranth. 6 or 8 medusa-buds, in various stages of develop-
ment, are arranged in a circlet situated upon some of the hydranths between the distal base
and the circlet of tentacles. The terminal hydranths and those near the free end of the main
stem bear no medusae, for these appear to be developed upon the old hydranths. The hydroid
of Nemopsis is thus a Bougainvillin, but the medusae arise from the sides of the polypites,
not from the stems.

When set free the medusa is flattened and folded so that the manubrium projects beyond
the velar opening. In about an hour, however, it expands and begins to swim. It is then
about 0.5 mm. in height, and the bell is globular, being about as broad as high. The manu-
brium is short and tubular and without oral tentacles. In some individuals there are 8 ten-
tacles, 2 at the base of each radial tube. In others there are but 4 primary tentacles, one
upon each tentacle-bulb at the base of the 4 radial tubes.

As development proceeds the marginal tentacles increase in number, those which are
nearest the middle of the basal bulbs being the oldest. The new tentacles appear at the sides
near the base of the cone-shaped basal bulb. The 4 oral tentacles develop upon the 4 radial
sides of the manubrium, and finally the gonads begin to develop upon the 4 radial-canals
adjacent to the sides of the manubrium and finally extend down almost the entire length of
the radial-canals.

According to Hargitt, 1901, about 5 per cent of the medusae of TV. bachei display vari-
ations of a numerical nature in the radial-canals, manubrium, gonads, and tentacles.

Ncmopsis crucifera Haeckel.

Hipprxrcnr crucijera, FORBES, 1853, Trans. Roy. Soc. Edinburgh, vol. 20, p. 313, plate 10, fig. 5.
.\rertif}fnn cruriftra, HAECKEL, 1880, Syst. der Medusen, p. 635.

This species is distinguished from TV. bachcl by its dark-red color and possibly by its
more nearly spherical bell.

Bell oval, 4 mm. wide, 4 mm. high. Each of the 4 marginal tentacle-bulbs bears a median
pair of short, stiff, club-shaped tentacles, flanked on either side by 2 to 3 longer filiform ten-
tacles. There are thus 6 to 8 tentacles upon each marginal bulb. The stomach is cruciform,
and its sides extend half-way down the 4 radial-canals. The oral tentacles are short and branch
twice dichotomously. The gonads are as in TV. bachei. Forbes shows the stomach, gonads,
and tentacle-bulbs of a darker red color than has been observed in TV. bachei. The tentacular
ocelli are black. Found by Forbes and Haeckel at Tobermory and Handa Island, off the
west coast of Scotland.

AXTIlnMl-.lil ;,.]; KATIIKKA.

17.')

Genus RATHKEA Brandt, 1837, sensu Vanhbffen.

««//;icfl, BRANDT, 1837, Bull. Sci.Acad.St.Pftenbouig, nd. i.|>. 1X7; iX;X, Mom. Acatl.Inip.St.Prirn.bnurjvrr.fi. fine 4.,,.-;-.

rv/<r/*, in part, SARS, 1835, Beskriv. og Jagttag, p. 28.

l.izzi.i, FORBES, 1846, Annals anil Mag. Nat. Hist., ser. I, vol. 18, p. 286; 1848, British Naki-d-i-u-d Mrlusa-, p. 64.--

CLAPAREDE, 1860, Zeit. fur wissen.Zool., Bd. 10, p. 401, (af. 32, fign. 1-3.— AOASSIZ, L., 1862, C.me. Vie. HI-.I. I . s.,

vol. 4, |>. 345.— AC;ASSIZ, A., 1865, North Amer. Acal., p. l6l.---\Vv.\t R, ]XS<;, W,rhrl|..M-n .!.-> \V.-i- i-n \|..-r.--. p.

73.— GARSTANI., 1904, Journ. Marine Biol. Association, Plymouth, New Series, vol. -,, p. 213. \U^. iSir. M> m. M'i

Conip. Zool. at Harvard College, vol. 23, p. 14.— HARI.ITT, 1904, Bull. tr. S. Bureau "I 1 i li, n, . \..|. -4. p. 37.
Rtahlaa+tfargelllam+Laaaa,Cuu«, tXne, Biblmtheta Znnlngica, Hrft 19, l.fg. i, pp. 13, if>, 24, 3/1, tic.
Marfrllium, BRUWNE, 1X95, Proc. and Trans. Liverpool Biol. Sot., \ol. </, p. 270; 1896, Proc. Zooi. S,,( . London. Marftllium,

p. 477; /.iraia, p. 475.

Li*u'a+ Liailla+ Margellium-t- Rathkea, HAX-KM, 1X79, Svst. di-r Mi-du .-n, pp. 72, 82, 83, 94, 96.
Ralhkfii, LITKF.N, 1850, Vidensk. Mrddel. Nat. r..n-n. Kl"l"-nh.ivn. pp. iX, • <;. A..ASMZ, I,., 1X1,2. Cone. Vie. Hit. 1 . s.. vol.

4. p. 541;. -MtTsc-iiMKOKt, 1886, Embryol. Studien an Mrdu&rn, pp. 23, 37, 48, etc.— VANIIOI i i\. 1X91, /.,„.!. An/. il;. i .

[ahrg. 14, p. 445.
Liz:ia± RofMea, MAA», e 897, Mem. Museum Comp.Zool.at Harvard C'olli-p'. vol. 23. p. 14; njo5,Cr.isprdotrn Mednsrn dcr Xihaga

Expedition, Monoj;. 10, p. 12. -Bioor, 1905, Revue Suissc dc Zool., tonic 13, pp. e?7, 144 (illation of references to 1850).

The type species is Ratliki-n t>l<i>n,-iilin,hii Brandt, of the Black Sea. 1 believe future
studies will show that this medusa is identical with Rntlikr/i rjctnpiiiiitntn.

In 1846-48 Forbes introduced the generic name " Lizziti," applying it to I.IT.ZHI mtn-
piiiictntti, which Browne and Garstang have shown to be identical with ('. \ttm in /o/>j//i, tutu
of Sars, 1835.

Haeckel, 1879, defines the following genera:

Haeckel

's name.

Lizzia.

Margelhum.

Lizzella.

Rathkca.

Condition of oral tentacles.

Condition of the Xclustersof
marginal tentacles.

Unbranched .

4 radial clusters have
more tentacles than
the 4 interradial.

Branched.

4 radial clusters have
more tentacles than
the 4 interradial.

I'nbranchcd.

All X tentacular
clusters alike.

Branched.

All S tentacular
clusters alike.

Vanhiiffen, 1891, shows that " Lizzia, Margellium, and Ltzzflla" are stages in the growth
of Ruthkca.

Much confusion has thus been introduced by the too narrow definition of genera, as
restricted by Haeckel, and it seems well to broaden the genus so as to make it apply to these
medusae in all stages of their free lite.

GENERIC CHARACTERS.

Margelinae with simple or branched oral tentacles and 8 clusters of marginal tentacles,
4 radial and 4 interradial.

There are perradial, adradial, or interradial gonads on the sides of the stomach and the
oral tentacles terminate in knob-like clusters of nematocysts.

In Rat/ikca medusa-buds are commonly developed upon the sides of the manubiium.
Chun, 1895, has shown that these buds are produced entirely from the ectoderm of the parent
medusa. A short time before the young medusa is set free, however, a connection is estab-
lished between the gastrovascular cavity of the bud and that of the mother medusa. Claparede,
1860, describes a "Lizzia" from Scotland in which the embryos are developed upon the outer
walls of the stomach very much as is the case with the planulae ot Brmgantvillia superciliaris.
In the case of this "Lizzia," however, the eggs, or embryos, are set free inclosed in capsules
and then develop directly into medusae. This remarkable observation awaits confirmation.

According to Browne, 1896 (Proc. Zool. Soc. London, p. 477), the medusa of Lizzm ( Rut liken )
nftnpunftata first reproduces itself by means of medusa-buds upon the_manubnum, and when it
reaches the adult condition either ova or spermatozoa develop within the ectodermal wall of
the stomach, the sexes being separate; and the same statement applies to Rntlik,-<i hl'inJnifi.

It is highly probable that the very variable R. oi-topunfttitn is found in the Mediterra-
nean and along the Atlantic coasts of Europe and America, and that R. blumenbachii and
R. hlnnJiiin are onl snonyms. R. fn.\,-i,-nliil<i is the only known Anthumedusa in which

the gonads are restricted to the

sides of the maiuibiium.

176

MKDUS.E OF THE WORLD.

Tabular Synnpsis of the Species of Rathkea.

Rathkea blumen-

R. formosissi-

R.octopunctata

R.octopunc-

R. fasciculata =

R.octonemalu

R.elegans

R.blon-

bachii Brandt,

ma= Lizzia

= Cytaeisocto-

tata var.

Melicerta fasci-

Maas, 1905.

= Lizzia

dina =

1837 to 1838.

formosissima

punctata Sars,

grata = Liz-

culata Prron et

elegans

Lizzia

Browne.

1835 to 1846 =

zia grata A.

Lesueur. Liz-

Mayer,

blondina

Lizzia octo-

Agassiz, 1862

zia kollikeri

1900.

Forbes.

punctata

to 1865.

Gegenbaur,

Forbes, 1848.

i8<;4 to 1856.

Shape

Bell-shaped.

Bell-shaped.

Pvriforrn. 5 high,

Pyriform.

Globular.

Globular,

Apex

Pyriform .

and

6 high, 4 wide.

3 high, 2.5

4.5 wide.

3 to 5 high, 3

Widest at up-

dome-like.

dome-

6 high, 4

size of

wide.

wide.

per two-thirds

6 high, 5

like, sides

wide.

bell in

of bell. 1 5 high,

wide.

vertical.

mm.

12 wide.

6 high, 5

wide.

No. of

3 in each cluster

5 in each ra-

4 to 5 in each ra-

? in radial, 3 in

15 to 20 in each

7 to 9 in

4 in radial,

3 in each

tenta-

(24 in all).

dial, 3 in each

dial cluster. 3 in

interradial

of the 8 clusters

radial, 5 to

3 in inter-

radia!

cles in

interradial

each interradial

clusters. As

when mature.

7 in interra-

radial

cluster. 4

each of

cluster.

cluster. When

in R.octo-

When only 3

dial clusters.

clusters.

small iso-

the 8

half-grown there

punctata.

mm. high there

lated in-

margin-

are 3 tentacles

are 7 in radial

terradial

al clus-

in each of the 8

and <; in inter-

tentacles.

ters.

clusters.

radial clusters.

Condi-

4 oral tentacles,

4 oral tentacles,

4 forked oral ten-

As in R. octo-

Each of 4 oral

Each of 4 oral

Branched,

4 simple,

tion of

each with 3 pairs

each with a

tacles, each

punctata.

tentacles,

tentacles

dichoto-

unbranch-

oral

of simple lateral

simple ter-

with pair of

branches dichot-

branches

mously

ed.

ten-

branches.

minal cluster

nematocyst-

omously 6 to 8

dichoto-

3to4

tacles.

and 3 to 5

knobs at its

times.

mously 4 to

times.

pairs of nem-

base.

5 times.

atocysts.

Form

A short, conical

Peduncle

Peduncle about as

Peduncle i .5

Peduncle not

Peduncle

AsinR.

As in R.

and

peduncle about

about as long

long as stomach.

times as long

quite as long as

about as long

octone-

octopunc-

size of

half as long as

as stomach.

Both together

as stomach.

stomach.

as stomach.

malis.

tata.

manu-

stomach. Both

about half as

Both to-

Both together

brium.

together half to

long as depth of

gether about

about half as

two-thirds as

bell-cavity.

two-thirds as

long as

long as depth

long as depth

depth of bell-

of bell-cavity.

of bell-cavitv-

cavity.

Gonads.

8 ( ?)interradial.

4 interradial.

When young.

As in R. octo-

4 perradial gon-

4 interradial

Interradial.

As in R.

No medusa-buds

When young,

medusa-buds

punctata.

ads, each gonad

horse-shoe-

No

octopunc-

observed.

medusa? pro-

developed in

forming a cres-

shaped. No

medusa-

tata.

duce medusa-

the 4 interradii

cent around

medusa-

buds.

buds.

of stomach.

place of en-

buds.

When older, this

trance of radial-

budding ceases

canal into

and sexual pro-

manubrium.

ducts develop.

Color

Tentacle-bulbs

?

Entoderm of

Entoderm of

Entoderm of

Brownish-red

Stomach

Stomach

browni sh-yellow .

stomach and

stomach and

stomach and

as in R. fasci-

bright-

and ten-

Stomach straw-

tentacle-bulbs

tentacle-

tentacle-bulbs

culata.

green,

tacle-

colored. Gonads

ranges from

bulbs ranges

red to brown-

tentacles

bulbs

and tentacles

brownish-green

from brown-

ish-red.

and canal

amber to

milky-white.

to black.

ish-green to

system

yellow.

orange.

pearly-

pink.

Where

Black Sea, Russia.

Falkland

Arctic Ocean and

Atlantic coast

Mediterranean.

Malay Archi-

Tortugas,

Coasts of

found.

Sebastopol,

Islands,

North Atlantic

of North

pelago.

Florida.

Great

April.

Atlantic coast

coasts of Europe

America,

Britain,

of South

and America.

north of New

France,

America.

Jersey.

and Ger-

many.

Distinct-

Distinguished by

Pinnate, oral

Oral tentacles

An American,

Dichotomous,

Very small

Brilliant

Simple, un-

ive

pinnate, oral ten-

tentacles,

forked, and pro-

local variety

oral tentacles.

basal stalks

green

branched

char-

tacles and short

and large

vided with later-

of O.octo-

Large number

of dichoto-

colora-

oral ten-

acters.

peduncle. These

number of

al nematocvst-

punctata.

of marginal

mouslv

tion.

tacles.

may be errone-

marginal ten-

knob; thus com-

tentacles.

branched

Dichoto-

ously described

tacles. Is

bining charac-

oral tentacles.

mous

by Rathke, and

this identical

ters of pinnate

Small num-

oral ten-

medusa is proba-

with R.octo-

and dichoto-

ber of mar-

tacles.

bly identical v ith

punctata ?

mous tentacles.

ginal tenta-

R.octopunctata.

cles.

ANTHOMEDrS.E — RATH K FA. 177

According to Gegenbaur, 1854 (Generationswechsel Polypen und Medusen, pp. 22 28,
Taf. 2, fign. I -9), the egg of Rathken fasciculata develops into a solitary polyp covered with a
horny peritheca and provided with 4 tentacles. The segmentation stages were studied by
Metschnikoff, 1886.

The oral tentacles in the young medusae are unbranched, hut in the majority of the
species they become torked as growth proceeds. In Rathken hlonJinn (I.izzia blonJinii
Forbes) they remain simple and unbranched throughout life. It appears best, however, not
to separate Lizzifi from the older genus Rathkea on the ground that I.izzia has simple tentacles
throughout its life, for all of these medusa; have simple oral tentacles when young and their
forked character in later life may better be considered specific rather than generic.

Rathkea blumenbachii Brandt.

Oceania blumenbachii, RATHKF, 1X55, Mem. Acad. Imp. Sci. St. Petersbourg, tomr 2, p. 321, laf. I, fign. 1-4.

(?)C\tifis octopunctata, SARS, 1835, Beskriv og Jagttag, p. 28, plate 6, figs. 14*2-14^.

Rathkia blurnenbachiana, BRANDT, 1837, Bull. Sci. Acad. St. Petersbourg, vol.i, No. 24, p. 187. Also: R. Oceania blumenbixhii,

1838, Mem. Acad. Imp. Sci. St. Petershourg, ser. 6, tnmc 4, p. 353.
Rathkea blumenbachii, HAFCKEL, 1879, Syst. tier Medusen, p. 96.
Rathkea blumenbachi, BF.DOT, 1905, Kevin- Suisse ile Zool., tome 13, p. 144 (literature- iS;i;-5o).

This medusa has a pyriform bell with thick, gelatinous walls. Bell-cavity about two-thirds
as deep as the bell-height. 8 marginal clusters, each with 3 tentacles all of equal length and as
long as the bell-height. 4 radial-canals. Stomach mounted on a short, conical peduncle.
Stomach and peduncle together are about one-halt to two-thirds as long as the depth of the
bell-cavity. 4 radially placed, oral tentacles, each of which forks at its end and gives rise also
to 4 short, side branches, i on each side of the tentacles. 8 gonads ( ?) No medusa-buds.
Stomach and tentacle-bulbs brownish-yellow. Sebastopol, Russia, in April.

For further description, see synoptic table of species of Rathkea.

Rathke's drawing is inaccurate in showing 8 radial-canals. The 4 interradial radiating
muscle strands were doubtless mistaken for radial-canals. I am inclined to suspect that this
medusa may be identical with the better-known Rathkea <it'tnpunctntu.

Rathkea formosissima.
I.izzia formosissima, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 278.

Umbrella bell-shaped, $ mm. high and 2.5 mm. broad, with a slight, transverse constriction
level with the top of the subumbrella cavity and a solid mass of |elly above it. 5 tentacles in
each radial, and 3 in each interradial, cluster. Stomach small, somewhat cubical, about as long
as broad, and situated on a broad peduncle about as long as the stomach. Mouth with a
plain, simple margin, somewhat quadrangular in shape. There are 4 oral tentacles, each with
7 to 1 1 clusters of nematocysts consisting in a terminal cluster and 6 to 10 lateral clusters in a
double row. Gonads on the stomach in 4 interradial, roundish swellings or masses. The young
medusa; have medusa-buds upon the stomach and only } tentacles in each group. Color! ?)
Found at Stanley Harbor, Falkland Islands, South Atlantic coast of South America.

Rathkea octopunctata Haeckel.

Plate 20, fig. 1 1 .

( >) Oceania blumenbachii, RATHKF, 1835. Mem. Acad. Imp. Sci. St. Pe'tersbourg, tome 2, p. 321, 4 figs.
Cyttris (?) octopunctala, SARS, 1835, Beskriv og Jagttag, p. 28, plate 6, figs. 14^-14^.

Cytais octopunctata, SARS, 1846, Fauna Littor. Noveg., tome I, p. 10, plate 4, figs. 7-ij.--l.iNKO, 1900, Travaux Soc. Imp.
des Nat. de St. Pctersbourg, tome 29, p. 152, fig. 2.

TT • . . ._»? . - O . » ,Hf_._XT_-U'. I _ .. O .

\taryellium octopunctatum+ Rathkea

17S MEDl'S.E OF THE WORLD.

Ralhkta oclopunclala, GIARD, 1888, Bull. Sci. France Belg., ser. 3, annee I, p. 317, plate 21.— CHUN, l896,Bronn's Thier-Reichs,
Bd. ^, Abth. l, p. 253, taf. 12 (development of medusa-buds).— BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 145
(all papers cited to 1850).

\largtllium actopunctalum, BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 15; 1905, Proc. Royal. Soc. Edinburgh, vol. 25,
p. 755.

Cubogasler gemmascens, HAECKEL, 1879, Syst. der Medusen, p. 76, taf. 6, figs. 8-[l (young budding medusa with only 2 mar-
ginal tentacles).

Adult medusa. — Bell pyriform,with solid, apical projection, and about 5 mm. in height and
4.5 mm. in diameter. Gelatinous substance thick at the apex of the bell, but thin upon sides of
bell-cavity. There are 8 clusters of marginal tentacles. 4 of these are radial and are situated
at the bases of the 4 radial-canals, and the 4 others are interradial and are situated 45° from the
radial clusters. In the fully-grown medusa there are usually about 4 or 5 tentacles in each
radial cluster, while the interradial clusters are each composed of not more than 3 tentacles.
When the medusa is not yet fully grown, however, there are normally 3 tentacles in both radial
and interradial clusters. Tentacle-bulbs large and swollen and filled with entodermal pigment-
granules. Velum well developed. There are 4 straight, narrow radial-canals. The manu-
brium is provided with a distinct peduncle which, however, does not extend more than one-
third of the distance from the inner apex of the bell-cavity to the velar opening. The gastric
portion of the manubrium is short and quadratic in cross-section. The mouth is surrounded
by 4 prominent lips. Each lip terminates in a pair of oral tentacles each of which ends in a
knob-shaped cluster of nematocysts. In addition to these there is also a pair of knob-shaped
clusters of nematocysts upon the sides ot each lip, centripetal to the terminal knobs. Medusa-
buds are developed upon the sides of the manubrium about the region ot the stomach. These
medusa-buds appear on interradially situated positions on the stomach-wall. According to
Chun (1895, Bibliotheca Zool., pp. 20, 21) the order of development of the medusa-buds

follows a very definite law. The oldest buds appear nearest
the peduncle of the manubrium and the youngest are found
9 lower down nearer the oral tentacles. The accompanying dia-

gram will serve to illustrate the law of succession in position
and in time ot the various buds upon the manubrium ( I ) being
the oldest bud and (16) the youngest (newest).

The oldest buds are developed near the proximal base of

the stomach, and the newest and youngest are found near the

bases ot the oral tentacles. Even before they are set free the

,0 young medusa? begin to develop buds on their stomach-walls,

'• and these secondary medusa-buds follow the same laws of

time and place ot development as do the primary medusa-buds

(see Chun, 1895, p. 23). Chun, 1895, showed that the medusa-buds were formed entirely from
the ectoderm ot the manubrium, the entoderm taking no part in their development. As the bud
becomes older, however, a connection is formed between its gastrovascular cavity and that of
the mother medusa. The developing medusa-bud has, at first, only 4 radial tentacles, one at
the base of each radial-canal. When set free, however, it possesses 16 tentacles: 4 clusters
each ot the 3 tentacles at the bases of the 4 radial-canals; and also 4 simple interradial ten-
tacles. The color of the entoderm of the manuhrium and tentacle-bulbs varies from brown-
green to black.

This medusa is common in the north Atlantic, where it has been taken off the coasts of
Norway, Barents Sea, Helgoland, England, Ireland, Scotland, and New England. Fewkes
found it in Narragansett Bay, Rhode Island, and it is common on the southern coast of New
England in March. Browne records this species from Valencia Harbor, coast of Ireland.
Levinsen, 1893, found it oflFthe west coast of Greenland. I believe that future studies will show
that R. blumenbachu of the Black Sea is identical with R. octopunctata. The drawing of the
oral tentacles of R. blumenbachu in Rathke's memoir is evidently diagrammatic and probably
quite erroneous. R. blumenbachu appears to be merely an R. octopunrtata in an intermediate
stage of growth wherein it has 3 tentacles in each of its 8 marginal clusters.

ANTHOMKIHS K KATIIKKA.

170

Rathkea octopunctata var. grata A. Agassiz.
Plate 18, fig. 5.

Lmia grata, AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 100, figs. 28, 29; 1865, North Amer. Acal., p. ihi,
figs. 251-258.— NUTTING, 1901, Bull. U.S. Fish Commission, vol. 19, p. 376, fig. 91 .— HARGITT, 1904, Bull. U.S. Bureau of
Fisheries, vol. 24, p. 39, plate i, fig. 4.

\oti l.Kzia grata, FEWKES, 1881, Bull. Mus. Comp. Zix.1. at Harvard College, vol. 8, No. 8, p. 142, platf I, figs. 1-7.

Margrllium gratum, HAFCKEL, 1879, Syst. der Mrdusen, p. 95.— LEVINSEN, 1893, Vid. Mi-ddrl. Nat. Fur. Kjnbenhavn (5), Bd .
4, p. 145.

Adult medusa (plate 1 8, fig. 5). — Bell about 3.5 mm. in height and 5 mm. in diameter;
pyriform in shape, and with well-developed apical projection which is solid. The gelatinous
substance is thick at the apex of the bell, but thin upon the sides. There are 8 clusters of mar-
ginal tentacles, 4 radial and 4 interradial. The radial clusters are each composed of 5 ten-
tacles, while the interradial clusters contain only 3 tentacles each. The tentacles are long and
flexible and are usually carried curled upward. The basal bulbs are large and filled with
entodermal pigment-granules. The velum is well developed. There are 4 slender, straight
radial-canals. The peduncle of the manubrium is long and extends about two-thirds of the
distance from the apex of the hell-cavity to the velar opening. The gastric portion of the
manubrium is small, and quadratic in cross-section. There are 4 prominent lips, each of which
terminates in a pair of oral tentacles which end in knob-shaped clusters of nematocyst-cells.
There is also found a pair of knob-shaped nematocystic clusters upon the sides of each lip.

Medusa-buds are developed in the interradii
upon the sides of the manubrium. The ento-
derm of the tentacle-bulbs and manubrium
is usually intense orange, but in some cases
the stomach is sage-green and the tentacle-
bulbs russet.

Young medusa. — In a medusa 1.2 mm.
in height there are 16 tentacles: 4 radial clus-
ters each consisting of 3 tentacles and 4 sim-
ple, interradial tentacles. The peduncle is
long and has the shape of the frustum of a
cone. The stomach is short and there are 4
simple lips which terminate each in a single
No medusa-buds are yet to be seen upon the

FIG. 93.— Rnlhken " kollikrri," = R. fasciculala, afti-r Crgrnbaur,
in Zeit. fiir wissen. Tool., Bd. 8.

knob-shaped cluster of nematocyst-cells.

manubrium.

I he medusa-buds upon the manubrium of the adult have at first only 4 simple tentacles,
one at the toot of each radial-canal. At this stage there are also 4 small interradial tentacle-
bulbs, which lack tentacles. It is not known what condition the medusa-buds may be in at the
time of their liberation from the parent.

This variety is probably an Arctic form. It appears in Newport Harbor, Rhode Island,
and in Massachusetts Bay during February, and disappears before June.

It is distinguished from the European and American /,. <n •tuftunitntn by its color and the
great length of the peduncle of the manubrium. Also the medusa-buds do not seem to make
their appearance until some time after the liberation of the young medusa, while in Lizziti
nit'jpuiictiitii they develop upon the stomach of the young medusa while it is yet attached to
its mother.

Rathkea fasciculata Haeckel.

la fasciruliita, PERON vr LJSCM'R, 1804, Ann.il. itu Mu-.. Hist. X.il.. I.

13, p. 14^ (.an literature to 1550;.
Lizzia kollikeri, GFGENBAUR, 1856, Zeit. fiir wissen. Zoo]., Bd. 8, p. 225, taf. 7, fign. ^ ij. K » > KM i is i \i> EMI > RS, 1862,

Zoolog. Beitrag, p. 84, taf. 13, fig. lo.
Bougaini-illia kotlikrri, LEUCKART, 1856, Archiv. fijr Naturgescli., Bd. 22, p. 24, taf. 2, fig. 2.

ISO

MEDUS.E OF THE WORLD.

This large Mediterranean form is described in the synoptic table of species of Rathkca.
Gegenbaur, i854(Generationswechsel und Fortplanz.bei Medusen, etc.. p. 22, taf. 2,fign. 1-9),
raised the larva until it developed into a solitary hydroid, 0.5 mm. long, with 4 tentacles,
each beset with a row of nematocysts. A horny perisarc covered the hydrorhiza.

The stages of segmentation were studied by Metschnikoff, 1886. The egg is o.ii mm.
wide and yellowish to white in color. The segmentation is total and equal, and results in the
formation of an elongated, ciliated, one-layered blastula with a large, central, segmentation
cavity. The ectoderm is formed by the cells which wander into the segmentation cavity by
ingression from the hinder end of the larva.

The medusa is quite common at Naples, Italy, during the winter. The bell is barrel-
shaped with a flatly-rounded apex. It is 8 mm. high and 9 mm. wide. The marginal ten-
tacles are highly contractile. There are 10 to 13 tentacles in each of the 8 clusters. Each

FK.. <)).-- Rtitlikea ftiscicitlalti. From living and preserved specimens, hy the author. Zoological Station, Naples, Italy.

A, Side view of medusa with tentacles strongly contracted. R, Tentacle showing ocellus. (', Ahoral view of
manubrium. /), View of oral tentacle, contracted.

tentacle bears an ocellus on its inner side. The oral tentacles branch dichotomously 7 times.
Whenever the medusa is disturbed hy a mechanical shock the oral tentacles suddenly expand
to their full extent, torming a network of filaments around the mouth, and thus it is seen that
through this fortuitous reaction the medusa captures its prey. Such a reaction is remarkable,
for in common with other medusae all other parts ot the animal contract when disturbed.

The 4 gonads form horseshoe-shaped, swollen crescents with transverse furrows. Each
crescent surrounds the point of entrance of the radial-canal into the stomach and the horns
of the crescent are directed ahorally. There are thus 4 pcrraJial gonads. This being the
only instance of perradial gonods seen in the anthomedusa?.

Rathkea octonemalis Maas.

Ralhkea aclonrmalis, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 12, taf. 2, fign. 1 1, 12.
( ? ?) Lizzflla octella, HAECKFL, 1879, Syst. der Medusen, p. 84.

Bell 5 to 6 mm. high, 4 to 5 mm. wide, with thick, gelatinous walls, bulging sides, and
flat top. There are 8 clusters of marginal tentacles. 7 to 9 tentacles in each ot the 4 radial,
and 5 to 7 tentacles in each of the 4 interradial clusters. These tentacles are not one-fourth
as long as the bell-height, and there is a black ectodermal ocellus upon the inner side of each
tentacle near the point of origin of the tentacle from the basal bulb.

The 4 radial-canals and the ring-canal are straight, narrow, and simple. The stomach
is mounted upon a short, wide, conical peduncle. The mouth is surrounded by 4 clusters of
oral tentacles. Each cluster arises from a short basal stalk and branches dichotomously
4 to 5 times. The basal stalks of these clusters are so short and the forked branches so long
that the mouth appears at first sight to be surrounded by 8 clusters of tentacles. This impres-
sion is, however, erroneous, for there are but 4 clusters of oral tentacles. The mouth does not

ANTHOMEDrS.-TC— RATHKKA.

1X1

extend to the level of the velar opening. The gonads are on the 4 interradial sides of the
stomach. They are doubly cleft in each interradius so as to present the appearance of 4
longitudinal hands below, but are (used above, forming a double horseshoe. The tentacle-
bulbs have decided red, entodermal pigment. The stomach is reddish-brown and the eggs
appear as flecks of yellow.

Found in the Malay Archipelago, Ternate, 3° 46' S. lat., 127° 47' long. E. from Greenwich.

It is possible that Lizzt-lln nitflla Haeckel may be identical with this species. Haeckel

gives a very brief description ot this medusa as follows : Bell egg-shaped, i 5 mm. high, 10 mm.

wide. Stomach globular or almost cubical, about
as long as its conical peduncle. Gonads 4-teathered
leaves in the stomach-wall. 8 simple, unbranched,
oral tentacles as long as the stomach. (Broken?)
8 marginal tentacle clusters, each with 8 long ten-
tacles. Coast of Japan. Color ( ?)

Rathkea elegans.

Plate 1 8, fig. 4.

Li-Kin elegant, MAYKR, 1900, Bull. Mus. Comp. Zool. at Harvard
College, vol. 37, p. 44, plate 38, fig. 127.

Bell 3 to 7 mm. in height and gelatinous
substance of moderate and nearly uniform thick-
ness. The sides of the bell are almost straight and
vertical and the top is dome-shaped. There are 8
groups ot marginal tentacles, 4 radial and 4 inter-
radial. Each radial group is composed of 4, and
each interradial ot 3 tentacles. The tentacles are
quite stiff and curve upward and are only about halt
as long as the bell-height. There is a small, dark-
FIC.. 95.- Rathkra octoiiemalis, after Maas, in Hydro- brown, ectodermal ocellus upon the under (velar)

medusen XihaSa Expedition. sjde Qf each tentac|e near the l)asaj |,ujb_ Jht.

velum is wide and provided with strong muscles. There are 4 straight, narrow radial-canals
and a simple circular vessel. There is a well-developed, conical peduncle above the stomach,
down which the radial-canals lead in their course to the gastric sac. The gastric part of the
manubrium is cruciform in cross-section and pear-shaped in general longitudinal contour.
The mouth is a simple opening without prominent lips. I he oral tentacles arise from 4 radial
sides of the stomach at a short distance above the mouth. Each tentacle branches dichoto-
mously 3 times, and then each tip terminates in 3 small branches which are covered with
nematocysts. The entodermal cells of the oral tentacles are chordate. The genital products
are situated upon the tour interradial sides ot the stomach.

The entoderm of the tentacle-bulbs and radial-canals is ot an intense, opaque, pearl--
white color, often displaying a tinge of pink. The entoderm of the stomach is of an intense
green, and the oral tentacles are pearly-pink. The intense opaque color of the radial-canals
contrasting with the hyaline transparency of the bell renders this medusa one of the most beau-
tiful found at the Tortugas, Florida. Several specimens were captured early in July, iS<yc;.

Rathkea blondina.

Lis

I.l
Dy

Liz

l>\
Dv

ia hlandina, FORBES, 1848, British Naked-eyed Medusa-, p. 67, plat.- 12, fig. 4.— BI.DOT, 1905, Revue SUISM- 7..JC.I., Bd. I •„
p. H7 (literature 1X48-50).— BROWNE, 1895, Trans. l,ivrrp<»>! Biol. Sue., vol. 9, p. 265; 1896, 1'r.n . /on]. Sor. of Lon-
don for 1896, p. 475 (review of recent literature, etc.); 1898, Journal M.ir ..... liiol ._ A •-. >. .. V S., vol. 5, p. rSX.
(FORBES), BROWNE, 1896, Proc. Zool. Soc. London, p. 475

ia, s/>., CLAPAREDE, 1860, Zeit. fur wissrn. Zool., Bd. 10, p. 401, taf. 32, fign. i-}.

marphesa minima (vounp stage)-!- Lizzia fltlparedei-^ L. blontlina, H*»rKFI, 1879, Syst. der Mr.luvrn. pp. -SI, 82, taf. 6,

fig- 7-
zia claparedei, CHUN, 1895, Bibliothi-ca Zoologies, Heft 19, p. 36 (budding of medusa-), figs. 2-4. HAK'KH.— Lizzia blonilina,

FORBES, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 754.
X,infd elhabetli,r, HAKCKKI., 1879, Svst. der Medusen, p. 83, laf. ft, fig. 12.
'inorphosa minima (young medusa), BROWNE, 1895, Trans. Liverpool Biul. Soc., vol. 9, p. 260.
smorphosa minima, HAECKEI.= Lizzia claparedei, HARTLAUB, 1897, Wissen. Meeresuntersuch. Deutsch. Meere, Ntue Fol

Abth. Helgoland, Bd. 2, p. 455.— HAECKFL= Lizzia clapandti, BROCII, 1905, Bergens Museum Aarbog, vol. u, p.

Foltv,
5.

IS'J MKnrs.E OF THF. WORLD.

Mature medusa. — Bell 6 mm. high, 4 mm. wide. Pyriform. There are 3 tentacles in
each of the 4 radial, marginal clusters and only one tentacle in each interradius. The stomach
is mounted upon a short peduncle, and both together are hardly half as long as the depth of
the bell-cavity. There are 4 radially placed, simple, unbranched, oral tentacles, each of which
terminates in a knob-like cluster of nematocysts and is as long as the stomach itself. When
immature the medusa produces medusa-buds upon the interradial sides of the stomach, but
later, when there are 3 tentacles in each radial cluster, the budding ceases and ova or sperm
develop. Claparede discovered that the eggs are inclosed in nematocyst-beanng capsules,
reminding one ot'the condition observed in Bougainvillia superctliaris. Browne states that the
medusa may become sexually mature when the bell is only 1.5 mm. high. The stomach,
gonads, and tentacle-bulbs are amber-yellow. Found off the coasts of England, Ireland,
Scotland, Germany, and Norway.

When the medusa is young there are only 8 marginal tentacles, 4 radial and 4 interradial.
Later there are 4 radial clusters, each consisting of 2 tentacles, and 4 smaller, isolated, inter-
radial tentacles. Medusa-buds are produced upon the interradial sides of the stomach, when
the medusa has only 8 tentacles. The 4 oral tentacles remain simple and unbranched through-
out life.

Lizzin eliztihftlicr Haeckel resembles R. blondina in all respects excepting that there are
4 tentacles in each radial, and 2 in each interradial cluster. Haeckel found this at Jersey,
English Channel. May it not be R. blonJina with more than the usual number of tentacles ?

Browne found a 3-rayed specimen of R. blonJina, with 3 radial-canals, j oral tentacles
} clusters of "perradial" tentacles each containing 2 tentacles and 3 single interradial tentacles

Genus CHIARELLA Maas, 1897.
C\iiaTflla, MAAS, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 15.

The type species and the only known form is Chiarella centripetalif Maas, from the
Gulf of California, Pacific coast of Mexico.

GENERIC CHARACTERS.

Margelina? in which the ring-canal gives rise to blindly ending, centripetal diverticula,
There are 8 cleft tentacle-clusters, so that the tentacles arise from 16 epaulet-shaped, marginal
swellings. 4 of these cleft tentacle-clusters are radial, and 4 interradial in position. The oral
tentacles are dichotomously branched.

This genus is chiefly distinguished by the blindly-ending diverticula of the marginal
ring-canal. The deeply-cleft tentacle-epaulets are quite characteristic, but this tendency
is also seen in some species of Bougainvillia and can not he considered as being a distin-
guishing characteristic of Chiarella, although it appears in a more marked degree in Chiarella
than in Bougainvillia.

Chiarella centripetalis Maas.

Chiarflla centripelalh, MAAS, 1897, Mem. Mus. Comp.Zool. at Harvard College, vol. 23, p. 15, taf. 2, fign. 1-4; 190^, Craspe-
doten Medusen der Siboga Expedition, Monog. 10, p. 13.

Bell high and dome-like, 15 to 20 mm. high, 10 to 15 mm. wide. Gelatinous substance
of slight consistency and fairly thick at apex, but thin at margin. Velum small. Bell-margin
exhibits 8 adradial, lappet-like emarginations which alternate with the radial and interradial
clusters of tentacles. The tentacles are very short, not more than one-tenth as long as the
bell-height and arising from 8 medianly cleft, marginal, epaulet-shaped swellings. The median
cleft in each tentacle-epaulet is so deep that the epaulet seems divided. 4 of these divided
epaulets are radial and 4 interradial in position. In the mature medusa there are somewhat
more than 40 tentacles in a single row on each double-epaulet, the medusa having altogether
about 320 tentacles. These tentacles arise side by side from the margin of the epaulets, and
the line of tentacles is continuous and not interrupted at the median cleft. There is an
ocellus upon the margin of the epaulet, on the velar side, at the base of each tentacle.

There are 4 straight-edged, narrow radial-canals and a ring-canal. This ring-canal gives
off a short, pointed, blindly ending, centripetal diverticulum in each interradius above the
median cleft of each interradial tentacle-epaulet.

AXTHOMKIM S.K --< HIAHKI.I.A.

183

The stomach is barrel-shaped and about one-fifth as wide as the bell-diameter. It is
not quite halt as long as the depth of the bell-cavity and is mounted upon a very short peduncle
down which the 4 radial-canals extend to the base of the stomach. There are 4 railiulh
placed clusters ot oral tentacles. Each cluster arises from a single stalk which branches
dichotomously 5 to 6 times, thus giving somewhat more than 32 terminal ramuli in each

quadrant. These terminal branches end in nematocyst-
knobs.

The gonads are 8 ad radial, linear swellings along the
sides ot the barrel-shaped stomach.

The entoderm ot the radial-canals and tentacle-bulbs
is rose-red, as is also the deep-lying entoderm of the
stomach. The gonads are dark-carmine and the ocelli
dark-blown.

Found in the (>ulf of California. Pacific coast of
Mexico, by the Albatross in the spring ot iSui.

Subfamily DENDROSTAURIN^, nov. subfam.

li '.thiilm' his ± ll'iUiail-ri MAAS, 1904, Sitzungsbcr. math.-plivs. K|.ISM- kgl.
Baver. Akad. Wissenschaft., Bit. 34, pp. 437, 458; Crjspeduten MI-I|UM-II
der tibogti Expedition, Monog. 10, pp. 15, 19. — Bigeluvs, nio<;, Mem.
Mus. Cninp. Zool. at Harvard College, vol. 37, p. 212.

C.tinnrjliil.e (in part), HAI:CKK|-, 187*), Syst. der Medusen, p. 140.

EH,. f)d.- -Clnnrella centripetalis, after M.I.I ,
in Mem. Museum Comp. Zoo],
at Harvard College.

SUBFAMILY CHARACTERS.

Dendrostaunnae: Anthomedusae with 4 or more pri-
mary radial-canals, some or all ot which branch. The
4 lips are without oral tentacles. The gonads are upon the interradial, or adradial, sides of
the stomach.

There are two tribes, Bythotiaridi and Williadi.

The tribe Bythotiaridi was established by Maas, 1904, as the family Bythotiaridae (Sit
/ungsber. inatli.-phys. Klasse kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 437; also, Cias-
pedoten Medusen der Si/won Expedition, Monog. 10, p. 15). Maas pointed out its relation-
ship to the Tiannae, which are exhibited most completely in the genus Bythotiara Ciiinther,
wherein the gonads are exactly similar to those ot the Tiarm;i', the only essential difference
being the branched radial-canals.

The tribe Williadi as we define it is equivalent to the family Williadae of Forbes, Agassiz,
Haeckel, Browne, Maas, and others.

In 1904, lor. fit., Maas showed that the Cannotidae of I laeckel, 1879, was a heterogeneous
group composed ot both Anthomedusae and Leptomedusx, and is, therefore, purely artificial
and must be abandoned.

\\ e will now proceed to define the tribes Bythotiaridi and Williadi and their »enera.

CHARACTERS ()!•• THE TRINE HYTHOTI ARUM.

Dendrostaurinx with branching radial-canals and a ring-canal. Without clusters of
nematocysts upon the exumhrella above the margin between the tentacles. With hollow ten-
tacles situated at the ends of the branches ot the radial-Canals. The <mnads are confined to
the inteiradial sides of the stomach.

1 he Bythotiaridi are separated trom the \\ illiadi by the presence ot a ring-canal and tin-
absence of clusters of nematocysts upon the exumbrella.

In the genera Bythotiara and Sihogitii the gonads are similar to those of the Amphine-
midi and I'and*idi of the Tiarinae, for they consist of ridges in the ectodeim of the inteiradial
sides of the stomach. Unfortunately the development ot these medusz is unknown, although
a peculiar process ot budding takes place in the tentacle-bulbs ol Niobia and the same medusa
is known to cast its eggs out into the water.

The genera of the tribe Bythotiaridi are as follows: Bythotiara Giimher, 1903; with 4
radial-canals which bifurcate so that 8 canals reach the circular vessel. S marginal tentacles,

1S4 MEnrs.E OF THE WORLD.

one at the end of each terminal branch of the radial-canals. Gonads in 4 interradial ridges on
the sides of the stomach. Manubrium with 4 simple lips. A single specimen found by Giinther
at a depth of 1,610 fathoms, N. lat. 52° 18.1', W. long. 15° 53.9'.

Sibogita Maas, 1904; with 4 principal radial-canals. Centripetal vessels arise from the
ring-canal and fuse with the 4 radial-canals. These canals are of various calibers. The
tentacles are hollow. The gonads are developed upon the 4 interradial sides of the manu-
brium in the form of a double row of transverse ridges in each quadrant. There are 4
simple lips.

S. geometries Maas, 1905 (Craspedoten Medusen der Siboga Expedition, Monog. 10,
p. 17, taf. 3, fign. 16-18.) A single specimen was found near the Celebes, Malay Archipelago.
S. simulans Bigelow, 1909, from the Eastern Pacific.

Niobia Mayer, 1900; with 4 main radial-canals, 2 of which bifurcate so that 6 radial-
canals reach the circular vessel. The 4 gonads are developed upon the interradial sides of the
stomach. The marginal tentacles develop into free-swimming medusae. The only species is
N. dendrotentacula of Tortugas, Florida.

Maas, 1904, is inclined to include the genera Netocertoides and Dichotomia among the
Bythotiaridi, but in these genera the gonads extend outward along the radial-canals and there
are solid interradial cirrus-like tentacles upon the bell-margin; they appear, therefore, to be
Leptomedusae.

CHARACTERS OF THE TRIBE WILUADI.

n'ilHad*, FORBES, 1848, British Naked-eyed Medusae, p. 19. — AGASSIZ, A., 1865, North Amer. Acal., p. 171. — HAECKEL, 1879,
Syst. der Medusen, p. I ^6. — BROWNE, 1904 Fauna and Geog. Maldive and Laccailive Archipelagoes, vol. 2, Pt. 3, p. 724. —
MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer, Akad. der Wissenschaften, Bd. 34, p. 43; 1905, Craspedoten
Medusen Siboga Expedition, p. 19. — HARTI.AUB, 1907, Nordisches Plankton, Nr. 12, p. 5.

Berenicida (sens, restr.), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 345.

H\<lt oluriilt, ALLMAN, 1871, Monog. Tubularian Hydroids. — DELAGE ET HEROUARD, 1901, Traite de Zool. Concrete, tome 2,
Par. 2, Les Coelenteres.

Dendrostaurinae with 4 or more primary radial-canals which, during the growth of the
medusa, give rise to one or more side branches. Stomach with 4 to 6 or more lobes upon the
interradial sides of which the gonads are situated. The tentacles are simple and hollow and
are situated at the ends of the main radial-canals and their branches. There are clusters of
nematocysts upon the exumbrella above the margin between the tentacles, and there is no ring-
canal. These last two characters serve to distinguish the- Williadi from the closely allied
tribe Bythotiaridi.

The tentacles are never grouped into clusters, but arise singly from the bell-margin. There
are neither cirri nor sensory clubs upon the bell-margin. The successive development of side
branches which arise from the main radial-canals is a distinguishing feature of this tribe.

It has been shown by Hincks. 1872, and Browne, 1896, that the medusa of Wtlhia stellata
Forbes develops by alternation of generations from the hydroid Lar sabellaruni of Gosse, 1857.

Brandt, 1838, and Browne 1904, found that there is no circular canal in Proboscidactyla
and Willsia, and in 1904 Browne showed that there is a tube under the clusters of nematocysts
of the exumbrella. Isolated nematocysts move along the bell-margin and then enter one of
these tubes and stop, and thus a cluster is formed by the accumulation of nematocysts.

For the Williadi we adopt the classification of Browne and Maas in the sense in which
they define the Williadae. Thus:

Proboscidactyla, BRANDT, 1835. Williadi in which 4 primary radial-canals leave the stomach.
Willsiu, FORBES, 1846, with 6 or more primary radial-canals leaving the stomach. The hydroid of //'.
stellata is Lor sabellarum.

It appears from researches of Hincks 1872, Browne 1896, 1904, and Maas 1904, that the
Williadi belong among the Anthomedusae and not, as stated by Haeckel, among the Lepto-
medusae. For Browne, 1896 (Proc. Zool. Soc. London for 1896, p. 468), demonstrates that
the medusa of ft'ilhia stellata Forbes is derived from a hydroid which was first described by
Gosse, 1857, under the name of Lar sabellarum. The best description of Lar sabellarum has
however, been given by Hincks, 1872 (Ann. and Mag. Nat. Hist., ser. 4, vol. 10, p. 313, plate
i ij I. The polypites are of two sorts and arise from a creeping hydrorhiza which is found grow-

ANTHOMKDUS.K — mTHOTIAKA.

185

ing upon the tube of Sabella. The sterile feeding-polypites arc fusifoim and have lint 2 ten-
tacles, which arise from one side of the body. The reproductive polypites, or lilastostyles, are
mouthless and cylindrical in shape. Their free distal end terminates in a knob-like cluster of
nematocysts. Several medusa-buds arise in a cluster from the side of the blastostyle. The
medusae become free and develop into It'illsin st,-ll<itii of Forbes, 1848 (p. 19, plate i, fig. i).

Tribe BYTHOTIARIDI.
Genus BYTHOTIARA Gunther, 1903.

Bvrhotiiira, Gi'NTHER, 1903, Annals and Mag.Nat.Hist.| Ger. 7, vol. II, p-424. MAAS, 1905, C'r.i:-pedoten MrJu m ,li r .s/'/ot;.;
Eiped., Monog. 10, p. 18.

The type species is Bythoti/iru murrn\t Gunther, from a depth of 1,610 fathoms, off the
southwest coast of Ireland.

GENERIC CHARACTERS.

With 4 (to 5 ?) radial-canals some or all of which bifurcate so that S (to 10 ?) canals reach
the circular vessel at the margin. Tentacles at the extremities of the terminal branches of the
radial-canals. 4 mterradial gonads, in the form of swollen ridges on the sides of the stomach.
4 (or 5 ?) simple lips. No longitudinal nematocyst tracts upon the exumbrella.

Bythotiara murrayi Gunther.

R\tholiara murrayi, GiJNTHfR, 1903, Annals anil Mag. Nat. Hist., ser. 7, vol. 11, p. 424, plate 10, figs. 4. c.

Medusa 7 mm. in diameter. 8 long tentacles at the bases of the 8 terminal branches nt
the radial-canals. 4 radial-canals which bifurcate close to the manubrium, giving 8 terminal

97-

FHV. 97. — B\tholiara murra\i. After Gunther, Annals ami Mag. Nat. Hist.

FIG. 98.— Bvlltoliara murrayi. A. Side view. fl. Aboral view of manubrium. From .1 specimen in the
Puritan collection, No. 46, 1902; at the Stazione Zoologies, Naples. Drawn by the author.

branches. 4 lips. Gonads in 4 ridges along the interradial angles of the stomach. Color ( ?
A single specimen from a depth of 1,610 fathoms in N. lat. 52° 18.1', W. long. 15° 53.9', off
the southwest coast of Ireland. Dr. Lohianco of the Naples Station kindly permitted me to
study a specimen of this, or a closely allied medusa which was obtained by the Krupp yacht
Puritan from a depth of 1,000 fathoms near Capri, Bay of Naples, and is now preserved in
alcohol at the Naples Zoological Station (fig. 98). The bell is dome-shaped, 12 mm. high and
14 mm. wide and flattened laterally. The hell-margin is destroyed. 5 primary radial-canals
arise from the stomach and 4 of these bifurcate, the other canal being simple. Thus 9 canals
extend to the periphery of the bell. The stomach is 2.5 mm. long and there are 5 inter-

ISO

.MKDFS.E OF THE WOULD.

radial, swollen gonads, with transverse furrows and a longitudinal median groove. I believe
this to be identical with Giinther's medusa. Having 5 primary radial-canals it may be
abnormal.

Genus SIBOGITA Maas, 1905; sens, emend Bigelow, 1909.

Sibo^ii'i, MAAS, 1905, Craspedoten Medusrn Jrr Xibo^a Expedition, Monnp. 10, p. 16. -Bn^Lnw, H. B., 1909, Mem. Mus.
Cnmp. Znol. at Harvard Collegi-, vol. 37, p. 213.

The type species is Sibogita geometrica Maas, from the Malay Archipelago.

GENERIC CHARACTERS.

Bythotiaridi with 4 main radial-canals. The marginal ring-canal gives off a number
of centripetal canals which in young specimens end blindly, but in later stages they connect
with the cruciform base of the stomach. The gonads consist of a pair of ridges with cross
furrows on each of the 4 interradial sides of the stomach. The tentacles are hollow and are
situated at the ends of the larger canals.

It is probable that this genus has been phylogenetically derived from some Turris-like
medusa.

Sibogita geometrica Maas.
Sibogita geometric,!, MAAS, 1905, Craspcdoten Medusen der Siboga Eipcd., Monog. 10, p. 17, taf. 3, fign. 16-18.

Bell is 40 mm. high, 20 mm. wide; with bulging, laterally flattened, barrel-shaped sides
and flatly rounded top. Gelatinous substance uniform and quite thick. There are 16 slender,
hollow tentacles, with swollen, club-like outer ends. These tentacles arise from the ring-canal
at the bases of the 16 larger canals and they project from the sides of the bell at a short distance
above the margin, as in Narcomedusae. The entodermal cores of the tentacles extend through

the gelatinous substance of the bell to the ring-
canal, and the margin between the tentacles is
thrown into 16 gelatinous"lappets." Thetentacles
are about three-fourths as long as the bell-height,
but are highly contractile. Their bases are only
slightly swollen and there are no ocelli.

There is a marginal ring-canal which gives
rise to centripetal branches. According to Maas,
4 wide, main radial-canals arise from the 4 cor-
ners of the stomach and proceed straight to the
ring-canal. 2 or 3 alternately arranged branches
fuse with the sides of each of these main radial-
canals. The largest (oldest ?) branches are near-
est the corners of the stomach, while the branches
which fuse with the more peripheral parts of the
main canals are smaller as we proceed outward.
Altogether about 28 centripetal vessels arise from
the ring-canal. The 4 main radial-canals are of
large caliber, but the centripetal vessels are nar-
rower in accordance with their age, the latest
formed being narrowest. Dr. H. B. Bigelow has
shown that the "side branches " of Maas are actu-
ally centripetal vessels which arise from the ring-
canal and only later in life join the radial-
canals.

The stomach lacks a peduncle and is narrow
and about half as long as the depth of the bell-
cavity. It is 4-sided at its base and the 4 lips are

r n. . 99.— fiibogna geometrica, after Maas, in hrgrb. . ' . . _

Sibaga Exped., Monog. X. flaring with folded edges. There is a throat-tube

PLATE 19.

Fig. 1. Niolna dendrotentacula, young medusa immediately alter being set

free from the parent. Tortugas, Florida, May, 1899.
Fig. 2. Niobia dendrotentacula. Side view of medusa showing the manner

in which the tentacles develop into medusae.
Fig. 3. Niobia dendrotentacula. View of manubnum ot mature medusa

after the budding process has ceased and the ova become mature.
Fig. 4. Niobia dendrotentacula. Diagram showing the sequence in which

the medusa buds are set free from the parent medusa; " I " is set

free first, "7" last.
Fig. 5. Niobia dendrotentacula. Oral view showing the order in which

medusa buds are developed at the tentacle-bases. Tortugas,

Florida, May 25, 1899.

Drawn from life, by the author.

PLATE 19

r
--

.\\TIIOMK1HS.K - siltiKilTA, NlolilA. 187

above the mouth and the stomach is urn-shaped and cruciform in cross-section. The
4 gonads are on the interradial sides of the basal part of the stomach and consist of 2 bow-
like longitudinal ridges in each interradius. These ridges are crossed by furrows as in Ti.nin.i-.

The entodermal parts are brown or almost black, but this color may be due in some-
measure, il not whollv, to the method of prcsi-rvation.

A single preserved specimen is described by Maas. It was captured in an open vertical
net between 450 fathoms and the surface, in the Malav Archipelago, near the Celebes, in i ' 28'
N. lat., 124° 28.2' E. long.

Sibogita simulans Bigelow.

SiboRita simulans, Bn.now, H. B., 1909, Mem. Mus. Cmnp. Znnl. at Harv.ir.1 CcdliTi-. rol. ;-. p. ;i;. pl.it.- ;, 41. an, I 4;.

This medusa may prove to be identical with S. ^i-nnii-him Maas.

Bell 30 mm. high, 22 mm. wide, laterally flattened, apex dome-like and rounded, gelati-
nous substance thick. Manubrium barrel-shaped, one-third as long as depth of bcll-caviiy.
Its base is 4-sided. 4 interradial gonads each consisting of a double, transverse row of folds
on the sides of the stomach. Lips well developed. There are 4 main radial-canals. In young
medusae the ring-canal gives rise to 8 adradial, blindly ending centripetal vessels. In later
life these adradial vessels connect with the stomach and 4 new interradial vessels make their
appearance and also connect with the stomach. Thus the medusa has 4 radial-canals and
12 vessels which arise from the marginal ring-canal and secondarily connect with the cruci-
form arms of the stomach. When mature there are 50 tentacles, one opposite each canal
and the others somewhat irregularly arranged. The tentacles are long, hollow, and flexible
and terminate in a knob of nematocysts. They have no basal bulbs and no ocelli.

The gonads are reddish-brown; bell faint blue.

2 specimens were found on the surface in the tropical Pacific between the Galapagos
Islands and Panama; another from Behring Sea, 121 fathoms.

Genus NIOBIA Mayer, 1900.
Niohia, MAYF.R, 1900, Bull. Mus. Comp. Zool. at Harvard Gillci;<-, vnl . 37, p. ^(,.

GENERIC CHARACTERS.

Dendrostaurinae with 4 main radial-canals, 2 of which bifurcate, so that Ci canals reach
the circular vessel at the bell-margin. The gonads are developed on the 4 interradial sides of
the stomach. The mature genital products are found in the ectoderm. There are 4 simple
lips, but no oral tentacles. The marginal tentacles develop into medusa- by a peculiar process
ot budding combined with fortuitous growth and are set free into the water as independent
animals similar to the parent medusa.

The only known species is Niobia dendrotentacula of the Tortugas, Florida.

Niobia dendrotentacula Mayer.

Plate 19, figs. 1-5.
Niobia ilrndroltntarula, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 36, figs. 141-144, plates 42, 4-,.

Adult medusa (plate 19, fig. 5). — The bell is slightly flatter than a hemisphere and about
4 mm. in diameter. The outer surface is smooth and there is no apical projection. The
gelatinous substance of the bell is quite thin and uniform, but not very flexible.

4 radial-canals arise from the 4 corners of the stomach; but 2 of these canals, which are
diametrically opposite each other, bifurcate so that 6 radial-canals reach the circular vessel
at the margin, 60° apart.

There are 12 marginal tentacles, one at the foot ot each radial-canal and one inter-
mediate between each successive pair of radial-canals. These 12 tentacles are arranged in
a bilaterally symmetrical manner in accordance with age. The rtldest and the youngest
tentacles are situated at the ends of the 2 simple radial-canals and the remaining 10 tentacles
are arranged in bilateral symmetry in accordance with their various ages, the axis being in
the diameter of the 2 simple radial-canals and the oldest and youngest tentacles. Kach halt
of the margin is thus a reflection of the other, and the order in age of each tentacle is shown
in plate 19, fig. 4. where (l) represents the oldest, (2) the next oldest, and (/) the youngest

MKIirS.E OF THE WORLD.

and least-developed tentacles. Thus tentacles (i) and (7), the oldest and youngest, are
situated at the ends of the two simple radial-canals, while tentacles (2, 2) and (3, 3), the
second and third in order of age, are found at the hases of the two forked canals. In addition
to these there are the interradial sets of tentacles (4, 4), (5, 5) and (6, 6) in order of age;
and thus the medusa has 12 tentacles, 6 radial and 6 interradial.

It is remarkable that through a peculiar process of growth and budding each tentacle-
bulb is successively developed into a small medusa which resembles the adult and is set free
into the water. The oldest tentacle is the first to develop into a medusa and be set free, and
the others follow in the order of their age until all of the tentacles have been cast off. They
are immediately replaced, however, by new tentacles, but after every one of the original 12
tentacles has been developed into a new medusa, the process of forming medusae declines
and finally ceases, and then the parent medusa becomes sexually mature.

The first stage in the transformation of a tentacle-bulb into a new medusa is the appear-
ance of a hernia-like outgrowth upon the floor of the subumbrella close to the tentacle-bulb.
This hernia-like pro|ection, which is composed of both ectoderm and entoderm and has
its cavity connected with a radial-canal or the circular vessel of the medusa, is destined
to form the manubrium of the new medusa. Soon after it begins to develop 2 pointed out-
growths appear on the sides of the ad|acent tentacle-bulb and these soon develop into new
tentacles. These outgrowing tentacles become larger and soon a still younger pair appear
farther out on the side of the original tentacle-bulb and these are soon followed by another
pair of tentacles, between the original pair of side branches and the margin of the medusa.
Before this has taken place, however, 4 short canals (the bifurcated radial-canals of the
future medusa) develop and the circular canal completes its circuit by coalescence.

An opening then appears in the velum of the parent medusa immediately below the manu-
brium of the developing medusa and this constitutes the velar opening of the new animal.
The manubrium becomes cruciform in cross-section and finally the new medusa is con-
stricted off and is set free with 6 radial-canals, 5 well-developed and 6 small, immature ten-
tacles and a velum (plate 19, fig. i). Even before the young medusa is cast off a hernia-like
outgrowth has developed near the base of its oldest tentacle preparatory to a repetition of
the process of budding. In this remarkable process of budding we see that the simple radial-
canals, the nr.g-canal, the velum, and the oldest tentacle are stolen directly, so to speak, from
the parent medusa. The forked canals, manubrium, and younger tentacles are new growths,
and thus the bud is not compelled to develop all of its organs anew. The budded medusae are
very hardy when detached and swim actively about, and soon develop new medusae out
of their tentacle-bulbs.

The manubrium of the mature medusa is about as long as the depth of the bell-cavity.
There is an enlargement near the middle of its length; the 4 simple lips are well developed
and cruciform. The gonads occupy 4 interradial situations in the upper part of the ecto-
dermal wall of the stomach. After the budding medusae have been set free the gonads become
mature and the ova are large and project from the interradial surfaces of the stomach. They
are finally dehisced into the water. The entoderm of the manubrium, tentacle-bulbs, and
circular canal is ocher-yellow, all other parts being transparent.

This medusa was abundant at Tortugas, Florida, from May 21 to June 4, 1899; and a
few have been found early in June of every subsequent year. It is very active and thrives
well in confinement.

Tribe WILLIADI.
Genus PROBOSCIDACTYLA Brandt, 1838.

Proboscidactyla, BRANDT, 1838, Mem. Acad. Imp. St. Petersbourg, Sci. Nat., ser. 6, tome 4, p. 390.— BROWNE, 1904, Hydro-
medusz Maldive Islands, p. 725; 1901;, Cevlon Pearl Oyster Report No. 27, Medusa?, p. 135. — MAAS, 1904, Sitzungsbrr,
math.-phys. Klasse kg!. Bavrr. Akad. \\~issenschaften, Bd. 34, p. 438; 1905, Hydromedusen der Siboga Expedition,
p. 20 — BIGELOW, H. B., 1909, Mem. Museum Comp. Zool. at Harvard College, vol. 37, p. 218.

It'ilhta, McCRADV, 1857, Gymn. Charleston Harbor, p. 47.

It'illia, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 346.— AGASSIZ, A., 1865, North Amcr. Acal., p. 171 .— VERRILL, 1873,
Report Commiss. Fish and Fisheries for 1871-72, p. 735.

n\<r,mnola+ DicranocannaJrWilletta+ Proboicidaayln, HAECKEL, 1879, Syst. der Medusen, pp. 151, 156, 157, 159.

It'illia, FEVVKES, 1882, Bull. Mus. Comp. Zool. at Harvard College, vol. 9, No. 8, p. 299; 1884, Mem. Mus. Comp. Zool. at
Harvard College, vol. 9, No. 3. plate <;, fig. 19.

ll'ilhiii, HUXLEY, 1891, Anatomy of Invcrtcbrated Animals, p. 120.

PLATE 20.

Figs. I, 2, and 3. Proboscidactyla ornata, young medusae. Agassiz Labora-
tory, Newport, Rhode Island, June, 1892.
Fig. 4. Proboscidactyla ornata, young medusa with tentacles expanded.

Agassiz Laboratory, Newport, Rhode Island, July 3, 1893.
Fig. 5. Part of a tentacle of the medusa shown in figure 4.
Fig. 6. Proboscidactyla ornata, young medusa having 8 terminal branches

to the radial-canals. Agassiz Laboratory, Newport, Rhode Island,

July, 1892.
Figs. 7 and 8. Proboscidactyla ornata, young medusae showing variations in

color. Agassiz Laboratory, Newport, Rhode Island, July 26,

1895.
Fig. 9. Proboscidactyla ornata, mature female in which the radial-canals

have 12 terminal branches. Agassiz Laboratory, Newport, Rhode

Island, September 17, 1896.
Fig. IO. Proboscidactyla ornata. Oral view of a mature female in which the

radial-canals have 16 terminal branches. Agassiz Laboratory,

Newport, Rhode Island, September 12, 1892.
Fig. II. Rathkca octofiunctata. Half-grown budding medusa. Abundant

among masses of floating ice in the harbor. Woods Hole,

Massachusetts, March 4, 1907.

See page 178 for description of figure 11.
Drawn from life, by the author.

PLATE 20

1 1

5

- a -

10

ANTHOMEDUS.E — PROBOSC1 DACTVI.A. ISO

The type species of this genus is Proboscidactyla flavifirrata Brandt, from the North
Pacific. Its 4 mam radial-canals branch more profusely than in any other species of the genus.

The American species, P. ornntn, was first described by McCrady under the name H'illsia
ornata from Charleston Harbor, South Carolina.

GENERIC CHARACTERS.

Proboscidactyla: Dendrostaurinae having 4 primary radial-canals which give rise to simple,
or branched, side branches during the growth of the medusa. The tentacles are simple and
hollow and are situated at the extremities of the radial-canals and their branches. Upon the
exumbrella there are clusters of nematocysts which alternate in position with the tentacles.
There are no marginal clubs or cirri and there is no circular canal. The gonads are upon
the 8 adradial sides of the stomach extending outward along the sides of the 4 radial-canals.

Haeckel, 1879, separates that which we here designate as Proboscidactyla into 4 genera
as follows:

l. Dicranocanna, in which the 4 radial-canals branch once, giving 8 radial vessels which reach the bell-margin.

2. Dyscannota, in which the 4 primary radial-canals give rise each to 1 side branches, and thus ll canals reach the

bell-margin.

3. H'lllfila, in which the 4 original radial-canals give rise each to 2 side branches, and the first-formed side branch uf

each quadrant itself gives rise to a side branch, so that 1 6 canals reach the bell-margin.

4. I'rohoscidactyla, in which the 4 canals that rise from the stomach branch complexly, and branches arise from the pri-

mary branches, etc., so that 32 or more radiating canals reach the bell-margin.

It is evident that medusae of the genus ProbosciJactyla may successively pass through
stages represented by the genera Dicranocanna, Dyscannota, and H'illettn.

The American, Atlantic Proboscidactyla or nut a is sexually mature in the " H'llletta" stage,
but its southern variety gemmifera gives rise to medusa-buds while in the "D\scannota" stage.

Psythia prolijera, Agassiz and Mayer, 1902, Mem. Museum Comp. Zool. at Harvard
College, vol. 26, p. 143, plate i, fig. 3, may be a young budding Proboscidactyla?

Proboscidactyla flavicirrata Brandt.

Proboscidactyla flaricirrata, BRANDT, 1854, Recueil Actes seances publiques Acad. Imp. Sci. St. Petersbourg, p. 28 (of the "Separ-
ate"); 1838, Mem. Acad. Imp. Sci. St. Petersbourg, Sci. Nat., ser. 6, tome 4, Par. 2, p. 390, plate 19.— AIIASSIZ, A., 1865,
North Amer. Acal., p. 173, figs. 280-282. -BROWNI:, 1904, Fauna and Geography Maldive and I.accadive Archipelagoes,
vol. 2, p. 725. — BKDOT, 1905, Revue Suisse de Zool., tome 13, p. 144 (all papers to 1850).

ProboiciJactyla fiaricirrata-^- P. brfvictrrata, HAECKEL, 1879, Syst. der Medusen, pp. 159, 160.

ProbotciJactyta brti'icirrata, MIIRHACH and SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 178.

Bell thick-walled and dome-like to globular. About 12 mm. wide and 10 mm. high.
4 main radial-canals arise from the stomach, but these fork near their points of origin and
each branch gives rise to side branches, from one side only and these again give off similarly
arranged branches so that about 54 to 70+ terminal branches reach the bell-margin. There
is a short tentacle at the end of each terminal branch of the radial-canals. There are clusters
of nematocysts on the exumbrella alternating with the tentacles. There is no ring-canal.
Stomach small, with 4 recurved lips. Cionads upon the sides of the 4 radial pouches of the
stomach. There are neither medusa-buds nor stolons.

Stomach, gonads, and tentacle-bulbs dull yellow. North Pacific, coast of Kamtschatka
and Puget Sound, Washington.

This species is distinguished by the profuse and peculiar branching of its radial-canals.

Proboscidactyla ornata Browne.
Plate 20, figs. 1-10.

H'illiia ornata, McCRADY, 1857, Gymn. Charleston Harbor, p. 47, plate 9, figs. 9-11.

H'illia ornata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. V, vol. 4, p. 34(1. AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol.
9, p. 96, figs. 20, 21 ; 1865, North Amer. Acal., p. 171, figs. 2740-279. — VERRILL, 1873, Report Commiss. Fish and Fisheries
U. S. for 1871-72, pp. 455, 735.— FEWCU, 1882, Bull. Mus. Cump. /mil. at HarvaYd College, vol. 9, p. 299, figs. 22, 23,
non fig. 24; 1884, Mem. Mus. Comp. Zool. at Harvard College, vol. 9, No. 3, plate 5, fig. 19. — MAVF.R, 1904, Mem.
Nat. Sci. Museum Brooklyn Inst. Arts and Sci., vol. l, No. l, p. 13, plate 2, fig. 12. — NUTTING, 1901, Bull. U. S. Fish
Commission for 1899, vol. 19, p. 377. — HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 40, plate i, fig. 5.

Dvscannota JvsJipleura, HAECLEL, 1879, Syst. der Medusen, p. 152.

H'illetta ornata, HAECKEL, 1879, Syst. der Medusen, p. 157.

ProboscvJactyla ornata, BROWNE, 1904, Hydromedusa? Maldive Islands, p. 726.- MAAS, 1904, Sitzungsber. math.-phys. Klasse
kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 439.

( ?) Dicranocanna furcitlata, HAECKEL, 1879, Syst. tier Medusen, p. 156.

I'll)

OF TI1K WOULD.

S,-\iitiHv-iiinlitn- nirJusa (plate 2O, figs. 9 and 10). — Bell about 5 mm. in diameter and
slightly higher than a hemisphere with Baring, hell-shaped sides. Gelatinous substance
thick and rigid. There are 16 marginal tentacles, one at the foot oi" each terminal branch
ot' the radial-canals. 4 of these tentacles are at the ends of the 4 primary radial-canals. 4
younger tentacles are at the extremities of the secondary, 4 still smaller tentacles at the ends
of the tertiary, and 4 more at the ends of the quarternary canals. There are thus 16 marginal
tentacles, 4 in each quadrant. The tentacle-bulbs are large and hollow and their entoderm
is densely pigmented. The shafts of the tentacles are very contractile and capable of great
elongation. They are covered with ring-like clusters o( nematocysts, which become apparent
only during periods of contraction. There are neither marginal clubs nor cirri, but midway
between the tentacles, immediately under the surface of the exumbrella, are 16 slender, centrip-
etal canals which alternate with the tentacular radii in position. One or more clusters of
nematocyst-cells are found upon the exumbrella immediately over each ot these canals, and
according to Browne, 1904, these nematocysts travel singly up the canals from the bell-margin
and become lodged at intervals, forming accumulations.

4 main radial-canals arise from the 4 radial corners of the stomach. Each of these
canals gives rise to 2 side branches and the longest and oldest of these side branches also gives
rise to a side branch. Thus there are 16 terminal branches each one of which ends in the
cavity of a tentacle-bulb. There is no ring-canal, but only a solid cord of entodermal cells
connecting the tentacle-bulbs one with another. The manubrium is flask-shaped and the
mouth is nearly at the level of the velar opening. There are 4 radially arranged, recurved
lips with crenated edges. The 4 gonads are developed upon the 4 interradial sides of the

Fir.. loo. — Diagrammatic oral views showing order of development of radial-canals in Prohosciiiacl\la ornala.
I, youngest stage; IV, mature condition.

stomach. In the female the mature ova are large and prominent. The entoderm of the
stomach and gonads is ocher-yellow or greenish-yellow, while the entoderm of the tentacle-
bulbs is crowded with brownish pigment granules. In the very young medusa, however, the
entoderm of the manubrium and tentacle-bulbs is often pink.

Toung nifJusa. — The hydroid of our American species has not been observed. In the
youngest medusae seen by me, the bell is about 0.7 mm. in diameter (plate 20, figs. 1-4).
The gelatinous substance is relatively thinner than it is in the adult. There are 4 equally
developed tentacles, capable of much expansion (fig. 4), but these are usually seen contracted
in captive specimens. These tentacles are situated at the bases of the 4 primary radial tubes.
4 slender, blindly-ending, centripetal canals arise from the margin midway between the 4
tentacles, and there is a cluster of nematocyst-cells upon the exumbrella immediately over
each canal. The manubrium is a simple, short tube with 4 well-developed, quadratic lips.
The condition of the medusa in this stage is represented in text-figure 100, I, which repre-
sents an oral view of the animal.

When the medusa is about 1.25 mm. in diameter ( plate 20, fig. 6), a side branch arises
from each of the 4 primitive radial-canals, and at the same time a second set of tentacles
(2222, text-figure 100, 11) make their appearance. These new tentacles are about 22° 30'
away from the primary tentacles (iin). If one face the oral surface of the medusa, as is
shown in text-fig. 100, it will be seen that the secondary tentacles (2222) lie in positions
opposite to that of the hands of a watch relatively to the primary tentacles.

The third set of radial-canals (3333, fig. 100, in) arise from the 4 primary canals beyond
the origins of canals 2222. This takes place when the medusa is about 2 mm. in diameter.

.\\TJIO\IL IMS K |'|;IP|:I.^< HiAiTM.A.

The fourth set of radial-canals (4444, fig. 100, iv) arise from the sides of the second set of
canals. At the completion of this stage, the medusa has 16 canals and 16 marginal tentacles
which are about 22° 30' apart. This docs not occur until the medusa is about $.5 mm.
in diameter (plate 20, tig. 4) and some individuals appear to In-come sexually mature before
this stage is reached (plate 20, tig. j).

This medusa is common in Narragansett and Buzzard's Bays on the southern coast of
New England. It is rare, according to Brooks, at Beaufort, North Carolina. McCrady found
a single mature specimen and I have found several in Charleston 1 1 arbor. South Carolina, and in
the Bahamas and Tortugas. It has not been taken north of Cape Cod on the New Kngland
coast, but is found along the entire coast of North America from the Bahamas to southern
New England. Bigelow, 1909, has demonstrated that there is a closely related budding
variety in the tropical Pacific.

It is possible that " Dicranocanna furcillata," found by Ilaeckel off the Atlantic coast of
Morocco, may be identified with P. ornnta.

From South Carolina southward our I', rinnitii gives rise to a \arietv which produces
medusa-buds upon stolons at the 4 radial corners ot the stomach near the points of entrance
ot the 4 radial-canals, and Haeckel's medusa has stolons in these

Proboscidactyla ornata var. stolonifera Bigelow.

(?) Williia, s/i., HIDCLFV, 1877, Manual of AM.IMHI, nl Invertebrate, I Animals, London, p. 120, fig. 17.

(?) Proboscidaclyla tropica, BROWNE, 1904, Fauna and Geog. "I M.iUiu- ami I. ana, In.- \u hipeljjjoe-, \.,l. 2, part }, Hv,lr,>-

meilusii-, p. 727.
Preboscidactyla ftavtctrrtitfi Tar. aolonifera, MAAS, iijo^, Cr.ivpeilnten Me,kisen ,ler .ViV'^/j Kipe,liti<>n, p. 21, taf. 4, fign. 24-28;

1906, Revur Suij-st- ,le /no]., liinit- 14, p. 89.
Pryboscidaclyla omtita, vnr. stohniffiii, Bigelow, H. B., 11)09, Mt-tn Mus. Cnmp. X.uol. .it H.irvar,! College, vul. ^7, p. 220,

plates 6, 41.

Bell rounded, 5 mm. wide, 4 mm. high, with a low, dome-like, apical projection and
thin walls. 12 to 20 or more tentacles, one at the base of each terminal branch of the 4 main
radial-canals. These tentacles are about as long as the bell-radius and have large, spherical

basal bulbs which bear entodermal
pigment-granules. An equal number
ot meridional lines ot nematocysts
alternate with the tentacles. These au-
upon the exumhrella and extend trom
the margin about one-tilth to one-halt
the distance up the sides ot the bell.
There are a number ot clusters ot net-
tling cells upon each line. The axial
line is composed ot a core ot ento-
dermal cells which branch off trom the
solid cord ot entodermal cells which
constitute the degenerate "ring-canal."
The 4 mam radial-canals branch
quite irregularly after the second fork-
ing, so that 12 to 20, or more, canals
reach the margin. The general mode
ot forking is, however, similar to that
of /•". rjniiitti.

Medusa-bearing stolons develop
at the second and thud tm kings ot
each of the 4 radial-canals. There are thus 8 of these stolons, 2_in each quadrant. These
stolons do not all develop at one and the same time. The two oldest appear at the second
forks of the canals of one diameter, then the next two at the second folks ot the diameter
90° apart from the first, and then the 4 others follow in the same order at the third fork-
ings of the canals.

The stomach is wide, cruciform, and urn-shaped, as in other species ot the genus. I he
4 lips are recurved, and the manubrium is not longer than one-half to two-thuds the depth

Fin. toi .—Proboscidoclyla orntiin i-ar. stolonifrrti, after M.i. iv, in
Hr^t-h. Xihogit KxpeJ., Mnnog. X.

192 MEDUS.E OF THE WORLD.

of the bell-cavity. The gonads may be pale green or colorless. The entoderm of the ten-
tacle-bulbs is black, or pale green, or colorless. The gonads are upon the sides of the
stomach and extend even beyond the first bifurcations of the radial-canals. It is found in
the Malay Archipelago, and at Acapulco, Pacific coast of Mexico.

This is identical with the medusa found by Huxley in the Louisiade Islands, North
Pacific, and called P. tropica by Browne, 1904. In P. "• tropica" however, the medusa-
bearing stolons arise from the first branchings of the 4 main canals and not from the
second and third points of forking as in Maas' and Bigelow's form. Bigelow's studies have
shown, however, that the two are identical, P. tropica being only a young stage.

P. "tropica" has 16 terminal branches to its 4 main radial-canals. 16 short tentacles, 4
stolons at the points of origin of the first branches of the 4 main radial-canals. These
stolons are long and end in a knob of nematocysts. On one side the stolon gives rise to a
series of medusa-buds, the oldest being nearest the free lower end of the stolon. When about
to be set free the budding medusa has 4 simple, unbranched radial-canals and 4 tentacles.

Proboscidactyla ornata var. gemmifera.
Plate 21, figs. 1-3.

ll'illia ornata, BROOKS, 1880, American Naturalist, vol. 14, p. 670; 1881, Studies Johns Hopkins Univ. Marine Lab., vol. 2,

p. 144.

Il'illiit i>finmifera, FEWKES, 1882, Bull. Mus. Comp. Zoo], at Harvard College, vol. 9, p. 300, plate I, fig. 24.
Proboscidaci\la pemmifera, MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer. Akad. der Wissenschaften, Bd. 35, p. 439.—

BROWNE, 1904, Hydromedusa- Maldive Islands, p. 727.
l)\*< unriotti ^emmifera, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 47, plate 8, fig. 17.

This variety of P. ornata is peculiar in that it gives rise to medusa-buds which are borne
upon 4 stolons that arise from the radial corners of the stomach near the points of origin of the
4 main radial-canals. Several medusa-buds in various stages of development may usually be
seen upon each stolon. When set free, each medusa has 4 simple, unbranched radial-canals,
4 well-developed, radially situated tentacles, and 4 small, interradial tentacle-bulbs. There
are also 8 centripetal canals alternating with and intermediate in position between the tentacle-
bulbs and tentacles. There are no medusa-buds or stolons upon the medusae when set free
from the parent.

All of the budding specimens of Proboscidactyla which have thus far been observed were
in the Dyscannota stage, having 12 terminal branches to the radial-canal system and 12 mar-
ginal tentacles. Each of the 4 canals which leave the stomach gives rise to 2 side branches,
and thus there are 3 terminal canals in each quadrant; 12 in all. It is possible that after the
medusa has ceased budding the canals may increase in number so as to become 16. On the
other hand, P. ornata var. gemmifera, may never develop sex-cells or possess more than 12
canals. The common P. ornata, which lacks medusa-buds and has 16 tentacles and terminal
canals, is found wherever the budding variety has been observed, although the budding variety
is not known north of Beaufort, North Carolina, while the sexual form is abundant upon the
southern coast of New England. For many years it was believed that in this gemmifera form
there was constantly but a single cluster of nematocysts over each centripetal canal of the
exumbrella, between the tentacles, whereas in the sexual P. ornata there might be i, 2,
or occasionally 3, clusters over each canal. In 1903, however, I found two specimens of
the gemmifera form which had I, 2, or 3 clusters of nematocysts over each centripetal canal,
and therefore resembled the condition seen in the sexual P. ornata. No gonads have been
observed upon the manubnum of the gemmifera form and the stomach is therefore smaller
and more slender than in the sexual P. ornata.

P. ornata var. gemmifern is found from Beaufort, North Carolina, southward to the
Bahamas. On the whole its close similarity to P. ornata in the Dyscannota stage leads me to
believe that it is only an immature or budding variety of Proboscidactyla ornata.

Proboscidactyla "minima" Browne.

Proboscidatiyla minima, BROWNE, 1905, Pearl Oyster Fisheries, Gulf of Manaar, Suppl. Report, Roy. Soc. London, 27, p. 136,
plate 2, fig. 3.

Bell hemispherical ( ?),o.75 to I mm. wide, with fairly thin walls. 4 main radial-canals
with lateral branches. Number of lateral branches ( ?) 10 to 20 short tentacles 0.25 mm.
long with globular basal bulbs. Stomach with 4 longitudinal lobes with the gonads upon their
sides. Mouth with a sinuous margin. Clusters of nematocysts on the exumbrella near the

PLATE 21.

Fig. I. Proboscidactyla ornata var. gemmifera, showing a budding medusa
about to be set free. Tortugas, Florida, June, 1897.

Fig. 2. ProkosfiJtiftvln or tint a var. gemmifera, showing unusually dark
pigment in the tentacle-bulbs. Beaufort, North Carolina, No-
vember 21, 1904.

Fig. 3. Proboscidactyla ornnta var. gemmifera, showing small immature
medusa buds at the 4 radial angles of the stomach. Tortugas,
Florida, June 12, 1897.

Fig. 4. LaoJicea cruciatu ("calcarata"), young medusa. Agassiz Labora-
tory, Newport, Rhode Island.

Fit;. 5. Laodicea cruciatn ("calcarata"}, half-grown male medusa. Agassiz
Laboratory, Newport, Rhode Island, September 17, 1896.

Drawn from life, by the author.

PLATE 21

\\Tllo\II-.hrs.K l'lf<>li(>SMl>\<"n LA, WII.I.SU. I'.lo

margin between the tentacles, dull ot Manaar, neat Cevlon, in February. Color ( ?) Dis-
tinguished by its small si/e (when mature ?).

Proboscidactyla "occidentalis" Browne.

Il'illia oecideataJis, FMVK*S, 1889, Bull. Kssex Inst., Salem, vol. 21, No. 7, p. 109, plate 5, fig. 3.

Prabasriilarltla orciilrnlalis, BROWNF, 1904, Fauna and Geog. Maldive ami I.aciadive .-Wlnprlagors, vol. 2, p. 726.

Size ( ?) Bell pyriform, gelatinous substance thick. 4 main radial-canals, each ot which
gives rise to 2 side branches, which also branch; thus each main radial-canal reaches the mar-
gin bv 5 vessels. 2O tentacles with reddish basal bulbs. Gonads on the marginal sides ot the
stomach. 4 lips. A single cluster ot nematocyst-cells on the exumbrella between each suc-
cessive pair of tentacles. Island ot" Santa Cruz. California. Color (''} Is this only a young
stage ot P. flavicirrata :

Genus WILLSIA Forbes, 1846.

H'ilhi<it FORHFS, 1846, Ann. and Mag. Nat. Hist., ser. i, vul. 18, p. 268; |84S, British \aked-eved Mediis.e, p. it). (FIISSJ,

1853, Naturalist's Rambles Devonshire (.'oast, p. 359.
H'illia, AiiAssiz, I.., 1X62, Cont. Nat. Hist. U. S., vol. 4, p. 346.
ll'illtiii, PEACH, 1867, Journ. Roy. Institute, Cornwall, p. 357.
H'illin. HAH 'KFi , 1X79, Syst. der Medusen, p. li;S. BKOWNI, 1902, Annals Mat;. Vil. HIM., SIT. 7, Vol. 9, p. 280; 1904,

HvdnmiriUis.e Muldive Islands, p. 729. MAAS, 1904, Sitzungsher. malh.-pltvs. Klasst- kgl. Bayer. Akad. der WisttDSchaft.,

B.I. 34, p. 4V).

GENERIC CHARACTERS.

Williadi with h or more primary radial-canals, each ot which gives rise to one or more side
blanches all of which extend to the bell-margin.

The type species is tf illsni itfllntu Forbes, ot the northern coast ot Europe. Its hydroid
stock is I. /ir sabellarum Gosse.

Willsia stellata Forbes.

ll'ill^in stellata, FORRFS, 1846, Annals and Mag. Nat. Hist., ser. l, vol. 18, p. 268; 1848, British NakeJ-eve.l Medusa-,
p. 19, plate I, fig. i.— BF.DOT, 1905, Revue Suisse de Znul., tome 13, p. I <;2 (literature to 1X50). BRIIWM., n»O4, Kauna
anil Gcog. Maldive anil I.accadive Archipelagoes, vol. 2, p. 729.

ll'iiliti stelldln, HAKCKKL, 1879, Syst. der Meilusen, p. 158.

l.,ir suhfllMiim (hyilroul), Gossv, 1857, Trans. Lmnean Soc. London, vol. ;2, p. 113, plate 20.

Ltir s<ibell<irnrn, HINI ks, 1X72, Annals and Mag. Nat. Hist., ser. 4, vol. 10, p. 313, plate 19.

( ) lI'iHiu furrata, HAF.CKEI., Ibid., p. 158.

It'illsia slrllata, Forbes = Lar sabellarum Gosse, BROWNF,, 1X96, Proc. Zool. Soc. London, p. 4(18; 1897, Proc. Zool. Soc. Lon-
don, p. 818, 9 figs, (the best modern description).

Lar sabellariifii, BROWN*, 1905, Proc. Rov. Soc. Edinburgh, vol. 25, p. 753 (medusa only).

Bell dome-like, about g mm. wide and 8 mm. high, with thick gelatinous walls, evenh
rounded. Stomach 6-sided, with well-developed, lateral lobes upon the sides ot which the
jMinads are situated.

When the medusa is set free from the hydroid there are 6 straight, simple radial-canals,
60° apart, but when about I mm. wide a side branch grows out from each ot the () original
canals. These side branches grow out from near the middle points ot the original canals,
and the original canals then bend so that 12 vessels reach the bell-margin nearly 30° apart.
In the next stage another side branch arises, farther out than the first and on the opposite side
from each of the fi original canals, and finally a third branch grows out from the inner side of
the first branch. Thus each of the 6 original radial-canals gives rise to } branches, and hence
24 terminal canals reach the bell-margin. Owing to the bending of the radial-canals tin-
terminal branches are spaced with a fair degree of symmetry, but the intervals between the
ends are not constantly 15°, for the space between the main canal and the last-formed branch
is apt to remain wider than the other intervals. (See Browne 1905.)

There are 24 short tentacles, one at the end of each terminal branch of the radial-canals.
These tentacles have well-developed, basal bulbs and are tapering and shorter than the bell-
diameter. 24 short, blindly-ending, centripetal canals extend upward from bell-margin
under the surface of the exumbrella, alternating with the tentacles; and there is a cluster ot
nematocysts upon the exumbrella over each one of these centripetal vessels. There is no
ring-canal.

The d-sided stomach is short, not being longer than one-half to two-thirds the depth of
the bell-cavity and there are 6 simple recurved lips. I he stomach differs from the Amer-

194

MEDUSAE OF THE WORLD.

Radial-canals.

Tentacles.

Exumbrella
clusters of
nematocvsts.

I..ulifst stage.

6 unbranched.

6

6

Sri niul stage.

i side branch to each
canal.

12

12

Thin! stage.

2. opposite branches
to each canal.

IS

IS

Fourth antl adult
stage.

The first-formed side
branch branches.
Thus each main

H

24

canal gives off 3
branches.

ican If ' ills i a discovered by Professor Brooks, in being 6-rayed at its center, whereas the
center of the stomach of the American medusa is only 3-rayed.

The stomach and gonads are yellow to reddish-yellow and the tentacle-bulbs are dark
yellowish-brown or purple-brown. This medusa is found off the coasts of Great Britain and
Ireland. It is abundant at Valencia Island off the southern coast of Ireland.
A summary of its stages in growth may be presented thus:

The hydroid is " Lar sakellarum"
of Gosse and Hincks. It grows upon
the tube of Sabella. The polypites
arise at intervals from a creeping
hydrorhiza. The fusiform feeding
polypites are larger than the repro-
ductive ones and have each 2 ten-
tacles which both arise from one side
of the body. The reproductive poly-
pites have no tentacles and their free
upper ends are somewhat globular
and devoid of a mouth, although they
are armed with many nematocysts.
The medusa-buds arise in clusters of
3 to 4 on the upper sides of the body
of the polypite. When set free the bell is somewhat higher than a hemisphere and the
medusa has 6 radial-canals and 6 tentacles alternating with 6 clusters of nematocysts upon
the exumbrella. The stomach is 6-lobed and has 6 lips.

Haeckel, 1879, p. 158, describes a " ff'illia furcata" from St. Nazaire, Atlantic coast of
France. This may be identical with W. stcllata, but it is said to have the gonads isolated upon
the radial-canals. The lower edge of the stomach is said to be folded or crinkled and without
true lips. In other respects the medusa is evidently identical with W. stellata. I suspect this
of being merely a specimen of l¥ . stcllata in which the central part of the stomach has been lost
through accident and is in process of regeneration.

Willsia (?) varians.

Probosciilarlvla varians, BROWNE, 1902, Fauna ami Geography Maldive ami Laccadive Archipelagoes, vol. 2, p. 728, plate 54,
figs, i, 2.

Browne describes this medusa from a single specimen with an irregular, 6-lobed stomach.

The medusa was 3 mm. wide, 2 mm. high. 6 main radial-canals, each with I to 3 lateral
branches. 17 tentacles, one at the end of each terminal branch of the radial-canals. No
ring-canal. Clusters of nematocysts on the exumbrella. Stomach very irregular with 5 ( ?)
lips. No gonads, but with 2 medusa-buds upon 2 of the radial-canals near the stomach.
Tentacle-bulbs dark brown. Maldive Islands, Indian Ocean. This may he an abnormal
specimen of Probosctdactyla nrnata var. stolonifera.

Willsia mutabilis Browne.

li'illiti mutabilisj BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 280; 1904, Fauna antl Geog. Maldive and Lac-
cadive Archipelagoes, vol. 2, p. 729.

Bell slightly conical, 6 mm. high and 6 mm. wide. Margin of bell slightly inverted and
summit broad and round. There are 24 or more tentacles. 6 or 8 radial-canals arise from
the stomach and each gives rise to 3 or more branches. The stomach is small, with 6 or 8
lobes, and the margin of the mouth is complexly folded. The gonads surround the stomach
and its lobes. This species is very variable. Found by Vallentine at Stanley Harbor, Falk-
land Islands and briefly described by Browne. Color ( ?)

Willsia brooksii, sp. nov.

Beautiful drawings of a young stage and also of the adult condition of this medusa
were made by the late Prof. William K. Brooks, while he was at Beaufort, North Carolina,
and were found among his unpublished figures, after his death. They were kindly pre-

ANTHUMKIHS.E— WILLSIA.

L95

loia A ami B, Proboscidaclyia nrntn.i, r,u . grnitmffrfi showing .t \outi^ st-ip- witli vtnlnii, lic.iiuii; mr.in^.t [>u.ls.
I' lo K, viiiinR and rnaUirr st.igr nf H'illiia brooktii :,p. nm .; II.IVIMI; I. prinurv r.i.h.il-ran.ils, 24 lrrniiii.il r.iiniili-
Fnmi Beaufort, North C.irolina. Drawn from lift-, by 1'rofi-ssor Brooks.

196 MEDUS.E OF THE WORLD.

sented to me by the Department of Biology of fohns Hopkins University for publication in
this work, and it seems but fitting that the species should be named in honor ot the great
naturalist who discovered it. It is closely allied and possibly identical with the European
ffillsia stellata, although the 3-rayed center of the stomach appears to distinguish it.

In the young stage there are 6 simple, slender radial-canals, 60° apart. The beU-walls
are relatively thin and the bell somewhat higher than a hemisphere with a bluntly pointed
apex. The 6 tentacles are 5 to 6 times as long as the bell-diameter and have swollen,
nematocyst-bearing, outer extremities.

In the mature medusa the bell is flatter than a hemisphere, thick walled, with a shal-
low bell-cavity. Twenty-four tentacles alternate with 24 exumbrella, nematocyst tracts
each with several clusters of nettling cells. The manubrium has 6 lips. Stomach j-rayed
at center, but each ray forks, giving 6 ramuli, 6 primary radial-canals which bifurcate twice
giving 24 terminal branches. The gonads extend along the sides of the stomach. The
size and color can not be determined from Professor Brooks's drawings. Found at Beau-
fort, North Carolina. (See page 195.)

Order LEPTOMEDUSJE Haeckel, 1866.

Lefilomtduij-, HAECKEL, 1866, Gencrellen Morphologic, BJ. 2, p. Ivii; 1879, Syst. dor Meduscn, p. 1 1 1 .— BIGILOW, njo'j,
Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p. 147. — TORREY, 1909, Publications University California, Zool.,
vol. 6, p. 12.

CHARACTERS OF THE ORDER l.EPTOMEDUS^E.

Hydromedusae which arise by alternation of" generations from Campanularian hydroids.
The gonads are developed upon the radial-canals. When present the lithocysts are of ecto-
dermal origin.

We may distinguish three families as follows:

1. Thaumantiadae Gegenbaur, 1856. Without lithocysts, but with marginal sensory-

clubs or cordyli.

2. Eucopidx Gegenbaur, 1856. With lithocysts and with less than 8 radial-canals.

3. yEquoridae Eschscholtz, 1829. With lithocysts and with more than 8 radial-canals.

The Leptomedusae are probably descended from the more simply organized Antho-
medusae; this is shown by the fact that when first set free from the hydroid some of the Euco-
pidae, such as Phortis,lzck lithocysts, which develop later. Also the hydroid of Eutinn called
Campanopsis has nearly all of the characters of the Tubularian hydroids. Asexual production
of medusa-buds is extremely rare in the medusa? of the order Leptomedusae being known only
in Eucheilota paradox tea and Eirenc incJusifera. These medusae are therefore creatures of
the coasts and are rarely found far out at sea, for they can not maintain themselves in
situations unsuited to the growth ot their hydroids.

Family THAUMANTIADjE Gegenbaur, 1856.

Thaumantiada, GEGENBAUR, 1856, Zeit. fiir wissen. Zoo]., Bd. 8, pp. 218, 236, 268. — MAAS, 189}, Ergeb.der Plankton Exped.,
Bd. 2, K. c., pp. 64-65; 1897, Mem. Mus. Comp. Zool. at Harvard College., vol. 23, No. i, p. 19; 1905, Craspe-
doten Medusen der Siboga Expedition, Monog. 10, p. 23.

ThaumanliaJit+Cannolicif, HAECKEL, 1879, Syst. der Medusen, pp. 120, 140.

FAMILY CHARACTERS.

Leptomedusae without lithocysts. The gonads are developed upon or extend outward
over the radial-canals and are not confined exclusively within the walls of the stomach.

In so far as is at present known the Thaumantiadae develop through alternation of
generations from Campanularian hydroids. For our knowledge of the development of the
species constituting this family we are indebted to the researches of Wright, 1862, on Thau-
mantias; A. Agassiz, 1865, on Melicertum; Metschnikoff, 1886, on Laodicea; and Browne,
1900, on Dipleurosoma.

It is evident, indeed, that the Thaumantiadae are very closely related to the Eucopidae.
They resemble the Tubularian medusae (Anthomedusae), however, in the absence of lithocysts
and the prevalence of ectodermal ocelli upon the tentacle-bulbs. Indeed, all authorities agree
in considering the Thaumantiadae as more lowly organized than the Eucopidae and /Equorid;p.

LEPTUMEUL'S.E. 197

and probably nearly related to the forms from which the Campanularian medusae have
developed (see A. Agassiz, 1865, pp. 124, 125; O. and R. Hertwig, 1897, Nervensyst. und
Sinnesorgane der Medusen, p. 155; Haeckel, 1879, pp. 121, 125; Brooks, 1895, pp. 301-303).
Haeckel goes so far as to state that the medusae of his genus Tetranema (fhaumantias)
are the primitive forms from which the Campanularian medusa? (Leptomedusae) have been
developed!

Asexual budding of medusae upon the gonads is unknown in the Thaumantiadae.

Brooks, 1895, shows that the marginal sense-clubs of Landicea contain no tract- of concre-
tions. Their cores are entodermal and they arise from the level of the exumbrclla nerve-
ring, above the velum, and are identical in all their anatomical relations with the sense-clubs
of the Narcomedusae and Trachymedusae.

Haeckel, 1879, includes Gomonemus and the Williadi among the Thaumantiadae.
Gon ion emus is certainly not one of the Thaumantiadae, for it has marginal lithocysts; and
according to the researches of Hincks, 1872, and Browne, 1896, the Williadi are derived
from Tubularian hydroids and should be placed among the Anthomedusae.

The Thaumantiadae may be conveniently, although somewhat artificially divided into
three subfamilies:

1. Melicertinae, with simple, unb ranched radial-canals.

2. Polyorchinae, in which the radial-canals give rise to blindly-ending side branches,

and these side branches do not fuse with the circular vessel.

3. Berenicinae, in which the radial-canals give rise to side branches, and these side

branches connect with the circular vessel.

A description ot the genera of the Thaumanriadie follows:

SUBFAMILY MELICKRTINJE: With simple, unbranched radial-canals upon which the gonads arc developed. The lips are as

numerous as the radial-canals. Ocelli, marginal sense-clubs, cordyli, and cirri are often present. There are no oral tentacles.

Thaumantias ESCHSCHOLTZ, ifay^Tttranema+Thaumantias HAECKEL, 1879. 4 or more tentacles, 4 radial-canals,
and 4 simple lips. No marginal clubs or cirri. Hydroid: Thaumantias,

Laodicea LESSON, 1845= Qctenema+ Laodice HAECKEL, 1879. 4 or more tentacles, with marginal clubs or cirri or both
between tentacles. 4 radial-canals, 4 simple lips. Hvdrmd: Cuspidelln.

Mriirrrtum OK I;N, sfnsu AGASSIZ = Melicertellajr Mflirertum HAKCKKI, 1879. With 8 or more icnt.n lc-. S radial-
canals, 8 simple lips, ami without marginal sense-clubs and cirri. Hydroid: Mflicrrtum.

t\ffliffrtissti HAI.CKJL, 1^70. \\'iili S nr more tentacles. 8 radial-canals and X lips. With mariMn.il i , ,

cirri. This is equivalent t<i Melicertis$a± Mrlietrtidium HAECKEL, 1X70, p. 124.

Orrhi'-tonui HAKCKEI., 1879. More than 8 radial-canals and the same number of lips. With or without marginal
sense-clubs and cirri. Development unknown.

Timoides BlGELOW, 1904. 4 radial-canals. Ring-canal gives rise to blindly-ending, centripetal canals. Stomach upon
a peduncle. 4 gonads on the radial-canals. Numerous tentacles and cirri. Development unknown.

SUBFAMILY POLVORCHIN.T-.: 4 or more radial-canals which give rise to blindly-ending side branches that do nut reach the circular

vessel. Gonads upon radial-canals, or upon their side branches. Ocelli, marginal sense-clubs, and cirri are often pp-M-nt.

Staurodiscu3 HAECKKI , 1879. 4 radial-canals, each of which gives rise to 2 blindly-ending side branches. Gon.ids upon

4 mam radial-canals, also upon side branches. There arc marginal sense-clubs. Manubnum tubular, with 4 simple

lips. Development unknown.
Ptychogena A. AGASSIZ, 1865. 4 radial-canals, which give rise to numerous blindly-ending side branches upon which

gonads are developed. Manubnum wide and shallow. Mouth simple, large, cruciform opening. Development

unknown.
Polyorckis A. AC.ASSIZ, 1862-65. 4 radial-canals, which give rise to many blindly-ending side branches. A number of

free sack-shaped gonads arise from radial-canals. Bell-margin simple. Manuhnum long and tubular, and tin-p-
are 4 simple lips. The ring-canal is simple without centripetal branches.
S(ri/>f>sia, TORRKY, 1909. Similar to Pol \orch is, but with a gastric peduncle and without pinnate branches on the

radial-canals distal to the gonads. (See Appendix.)
Xpirorodon HAKCKEL, i^jy^Goneomeantlrus, KIRKPATRKK, 190}. Similar to /'O/VOT/IIJ, but ring-canal gives, rise to

blindly-ending centripetal branches and bell-margin divided into lappets. Development unknown.

SUBKAMILY BIRHWICIN.'}-;: Radial-canals branch and all branches connect with circular vessel. Lips as numerous as main
radial-canals. Ocelli, marginal clubs, and cirri often present.

Cannota HAFCKEL, 1879. 4 main radial-canals each of which gives rise in 2 simple side branches, and thus 12 canals
connect with the circular vessel. There are 12 gonads, I upon each terminal branch of the radial-canals.

Curifria PERON, 1807 = Btrcn ice HAECKKI,, 1879. The 4 mam radial-canals give rise to non-dichotomoua side
branches, and the side branches also branch. Gonads on the terminal branches of the radial-canals.

Dichotomia BROOKS, 1903. 4 mam radial-canals all of which divide dichntonmuslv twu nr more times. Tin-
extend outward from the sides of the stomach over the radial-canals and their branches.

DipUurosoma AXEL BOF.CK, 1866 = Tttracannota MAVKR, 1900. "\ or more main radial-canals, some or all of uhuh
give rise to non-dichotomous branches. Gonads on the canals adjacent to the manubnum. According to Browne,
the hydroid appears to be Cuspidella?

Toxorchis sens, emend., HAECKKL= Toxorcki$+ Cladotanna HAECKEL, 1X79. 4 or more main r.ult,il-t anal', u Inch
branch dichotomously one or more times. Tin- gonads are upon the outer branches -.f the radial-canals near tin-
circular canal.

Netocertoitki M\YKR, 1900. 8 main radial-canals, which branch dichotomously. The gonads extend outward from tin-
sides of the stomach along the radial-canals.

L98

MEDl'S.E OF THi; WOULD.

Genus THAUMANTIAS Eschscholtz, 1829.

Tliaumiintia* (in part), EsciisnioLTZ, 1829, Svst. der Acal., p. 102. — OKEN, 1835, Allgemeine Naturgesch., Bd. 5, p. 226.—
LESSON, 1843, Hist. Zooph. Acal., p. 176. — FORBES, 1848, British Naked-eyed Medusa, p. 45, etc.

Thaumaniia!. GI.C.KNBAUR, 1856, Zcit. fiir wissen. Zool., Bd. 8, p. 237. — WRK;HT, 1862, Quart. Journ. Microscop. Sci., N. S.,
vol. 2, pp. 221, 308. — HINCKS, 1868, British Hydroid Zoophytes, p. 178. — BROWNE, 1896, Proc. Zool. Soc. London,
pp. 480, 489. — HARTLAI B, 1905, Zoologische Jahrbucher, Supplement 6, p. 567.

Ten ii'iniiii 4 ThaumantieiSf HAECKEL, 1879, Syst. der Medusen, pp. 125, 127.

Halmomises, VON KENNEL, 1891, Sitzbcr. Nat. Ges. Dorpat, Bd. 9, p. 282.

The oldest species is "Thaumantias hemispheerica" Eschscholtz, 1829 = Medusa In-nu-
sfi/i.rru-a Gronovius, 1760; but Browne, 1896, shows that this is undoubtedly a Phialidium,
and has lithocysts which former observers had overlooked. "Thaumantias aeronaut ica"
Forbes, 1848, is probably a Pliinlitiiuni, and the type species appears to be Thaumantias
fdlularia Haeckel = LaoJicc ccllularia A. Agassiz, 1862, 1865.

GENERIC CHARACTERS.

Thaumantiadae with 4 gonads upon the 4 radial-canals. With 4 or more tentacles.
Manubrium without a peduncle and with 4 simple lips. No marginal sense-clubs or cirri.
The hydroid is Thaumantias.

Haeckel, 1879, would restrict Thaumantias to include medusae with 16 or more tentacles,
and he proposed a new genus Tetrancma for medusae with 4 tentacles. We adopt a broader
definition and restore the older interpretation of Forbes and others, and include medusae
with 4 or more tentacles under Thaumantias.

T/iaumantias is separated from Laodicea by the fact that there are no marginal sense-
clubs or cirri in Thaumantias, whereas these structures exist in Laodicea.

The hydroid of Thaumantias was first described by Wright, 1862 (Journ. Microscopical
Science, vol. 2, p. 221). The stem is simple (or branched?) and rooted by a thread-like
stolon; hydrothecae campanulate; polypites with a prominent funnel-shaped proboscis.
The stem is sometimes ringed throughout, sometimes only at the base and summit. The
calycles of the polypites are denticulate, the edge having about 7 teeth. There are 16 or
more tentacles in a single ring. Wright reared the hydroid from the eggs of Thaumantias
inconspicua Forbes.

Thaumantias is unfortunately a receptacle for all the medusae of Phialidium in which
lithocysts have not been observed. The older authors, previous to Gegenbaur's researches
published in 1856, generally failed to observe the lithocysts; hence our heritage of numerous
"species" of "Thaumantias." Browne, a most assiduous student of European medusae,
recently stated that he had never found a Thaumantias. I have never taken one in thousands
of surface-tows made along the Atlantic coast of the United States from Eastport, Maine, to
Tortugas, Florida, nor did I find a single specimen of Thaumantias in cruising over the

Pacific while serving as assistant to Dr. Alexander
Agassiz. Dr. Lobianco, however, kindly permitted
me to study some well-preserved specimens of a
medusa collected by him at Naples, which I believe
to be a Thaumantias.

Thaumantias eschscholtzii Haeckel.

Thaumantias esrlischoltzii, HAECKEL, 1879, Syst. der Medusen, p. 129,
taf. 8, fig. 4. — LEVINSEN, 1893, Vid. Meddel. Nat. Foran.,
Kjbbenhavn, ser. 5, Bd. 4, p. 145.

Bell flat and watch-glass-shaped, and 3 to 4
times as broad as high. About 10 to 12 mm. in
diameter. There are 240 to 280 slender tentacles
with globular basal bulbs which are provided with
ocelli. The tentacles are about half as long as the
radius of the bell. There are no sense-clubs or
cirri upon the bell-margin. The velum is well
developed. There are 4 straight, narrow radial-

rii,. 102.— Thaumantias esrlischoltzii, after

Harcki-l, 1879.

I.KI'Tu.Ml.m S.K THAI MANT1AS. 1111. 1

canals. I he manubrium is very shallow and there arc 4 prominent, crinkled lips. The 4
gonads are thick and cylindrical and occupy the middle regions of the 4 radial-canals. Kach
gonad is about halt as long as the radial-canal upon which it is situated. Coloi ' ?) This
medusa is found upon the western coast of Greenland.

Thaumantias rubrum.
Chromalonema filmim, K> «*>•,, iss^, Hull. Mus. Cmnp. Xool. .11 H.irwr.l l'o]|i-j;i-, \ol. .(. \,,. s. p. ',05, lit;. 41. I'l.iii- i.

Bell hemispherical and gelatinous substance thick. There are 12 to |f> short tentacles.
No lithocysts, marginal clubs, or cirri. There are 4 straight, radial tubes. Manubrium shoit
and there are 4 slightly crenulated lips. Gonads developed upon radial-canals adjacent tn
manubrium. Each gonad is about one-third as long as the radial tube upon which it is situated.
The ova are very conspicuous. The gonads and tentacles air i ed in alcoholic specimens.

Found oft the New England coast by the United Stales Fish Commission in i SSo and
iXXi. Were, the lithocysts destroyed by preservation in alcohol '. Si/.e i :)

Thaumantias cellularia Haeckel.

l.n'ldl'f ifilllLlf I/I, Af.ASSI/, A.. (Site;. Nnllll AllHT. Al.ll., ['. 12^. fit;*-. I<K. I'l1',

'nitiumiintitis telliiLii i,t, \\ \\ < kit, i S-tj, S\ >t . JIT M«-,|UM-M, p. 129. — Mi RBAI II .ill.t Siu AKh K, 1 90*, , \'T« . '/.>•• >\ . s,i, . I.. ,;
vul. I, p. 172, plate 17, fijjs. 2-26.

Hell thick, dome-like, about }O mm. wide, 20 mm. hiidi. About 100 tentacles \\ith wcll-
developed basal bulbs set closely one by the side ot the other. Tentacles about as Ion" as the
hell-diameter. Stomach very small and with 4 long, slender, folded lips, 5 to <i times as long as
the diameter of" the stomach. 4 folded linear gonads nearly as long as the 4 radial-canals
upon which they are developed. Stomach, gonads, and tentacle-bulbs \iolet.

Abundant oft the Pacific coast of Washington during the summer and autumn.

Thaumantias forbesii Haeckel.

\ail ThaUtlltUltiai lllfunj^fi'll, I'OKIUS, iSaS, British Xakr 1-cvi-d Mi'ilu^r, p. 41), plalr I I, t|t-. ^.

T/iiitini<iiitiii! forbeiii, HAM KM, 1X711, S\st. Jrr Mcduycn, p. 129.

Tttrancma eucopium (young or abortive medusa r), HAKIKU, Ihi<l., p. 125, t.tf. S, fi^. i, :.

Bell Hat, [6 mm. \vide, 5 to 8 mm. high, in tentacles longer than the bell-diameter with
large, globular basal bulbs, separated by wide intervals. Ocelli upon the tentacle-bulbs.
Stomach small, globular. Mouth with 4 large folded lips, 2 to 3 times as long as the diameter
ot the base ot the stomach. 4 ellipsoidal gonads upon the outer thirds of the 4 radial-canals,
touching the margin. Mouth and stomach light-violet. Gonads and tentacle-bulbs dark
violet. Ocelli black. Coast of" Norway, near Bergen, Haeckel, September, iSdg.

Tetranema ruivpttini Haeckel, 1879, appears to be the same medusa, only X mm. wide
\\ith only 4 tentacles, and with 4 folded, spindle-shaped gonads over the greater parts ot the
4 radial-canals. Found in the Straits of Gibraltar, by Haeckel, in March.

Thaumantias lacustris.

Halmiimiiti lacuslris, VON KKNNLI., 1891, Sit/her. Nat. Grs. Dorpat, B.I. 't, p. :S;.
Hiilirifjniisrs Itifuslrn, VON KSNNKI-, Ann. anil Mag. Nat. Hist., ser. 6, vol. 8, p. 259.

Bell about 2 to 2.5 mm. in diameter and slightly higher than a hemisphere. There are

|f> to 24 very long tentacles with well-developed basal bulbs, which contain a simple ring of
pigment upon their outer (centrifugal) sides. There are no marginal am. I he velum is well
developed, being broad and thin. The manubrium extends horn the inner apex of the bell-
cavity to the level of the velar opening. It is cruciform in cross-section, with a wide proximal
base. The mouth is simple, with 4 small, bluntly rounded lips. The 4 radial-canals are sinuous
for the first three-fourths of" their length from the 4 corners ot the stomach. In the last quarter
ot their length, near their point of juncture with the circular tube, they are sti aight and narrow .
The canals are broad in the sinuous region and here the gonads are situated. Kach gonad is
longer than the portion of the canal upon which it is situated, and it is therefore thrown into
frills and twisted sinuously. The bell is hyaline or fainth velhmish. The tentacles and bell-
margin are slightly milkv. The gonads aie jrellowish-brown.
if,

MKDU.S.E OF THK WOULD.

This medusa was discovered by von Kennel in a fresh-water lagoon on the east coast of
Trinidad, south of Mayaro Point, in a cocoanut plantation. The water in which it was
living was apparently perfectly fresh, not being salt to the taste. The lagoon was in commun-
ication with the sea during the rainy season.

Thaumantias maeotica Ostrooumoff.

Thaumantias rnxolicu, OSTROOUMOIT, 1896, Bull. Acad. Imperiale des Sci. St. Petersbourg, bcr. 5, tome 4, p. 401, plate i, figs.
2, 4. S-

Bell fuller than a hemisphere, with very thick gelatinous substance; 18 mm. wide, 10 mm.
high. 32 filiform, slender tentacles, twice as long as bell-diameter, with well-developed basal

bulbs. Dark-colored ectodermal ocellus on each
tentacle-bulb. Velum narrow. 4 straight, slen-
der radial-canals. Manubnum 4-sided, prisma-
tic, with 4 simple lips. About half as long as
depth of bell-cavity. No peduncle. 4 spindle-
shaped gonads on the middle thirds of the 4
radial-canals. Sea of Azov, Russia.

Dr. S. Lobianco of Naples kindly permitted
me to study some very well-preserved specimens
of a Thaumantias which he collected in the Bay
of Naples. At first I was inclined to regard these
as a new species, but now believe that they may
represent a young stage of Thaitmantias micotica.
I herewith present a cut (fig. 103) representing
the largest of Dr. Lobianco's specimens together
with the following description:

Bell 4 mm. high and of about an equal width.
Apex evenly rounded and exumbrella smooth.
Gelatinous substance thick at apex, but thin at
bell-margin. Bell-cavity about two-thirds as
deep as the height of the bell.

There are not more than 16 tentacles; the
number among six specimens observed by me
ranged from 8 to 12. The 4 perradial tentacles
are about twice as long as the bell-diameter, the
4 interradial about half this length, and the
adradial tentacles are not more than one-third
as long as the interradial. All of the tentacles
taper gradually to their tips and are sharply set
off from their globular, swollen, basal bulbs.
There is a black, abaxial, ectodermal ocellus
upon each basal bulb. The shaft of each ten-
tacle is regularly besprinkled with small, isolated,
wart-like patches of nematocyst-cells, which do
not form rings.

Velum very wide. There are 4 straight-
edged, narrow radial-canals and a simple, slender
ring-canal. Manubnum small, flask-shaped, and

I'll.. 103. — Thaumantias

with 4 simple lips. It is not more than one-fourth as long as the depth of the bell-cavity.
The 4 spindle-shaped gonads are developed along the 4 radial-canals from their proximal
thirds to their middle points. They are apparently immature.

The collection of the Stazione Zoologica at Naples, Italy, contains six well-preserved,
alcoholic specimens of this medusa, which were gathered by Dr. Lobianco in the Bay of Naples.

This medusa bears some resemblance to Thaumantias quadrata Forbes, 1848, British
Naked-eyed Medusae, p. 43, plate 9, fig. 2; from Loc Fyne, Scotland. In this British medusae,
however, there are apparently no true ocelli, the so-called ocelli of Forbes being orange-
colored entodermal pigment in the tentacle-bulbs.

PLATE 22.

Fig. i. Disi'jiifinti tiirriJa, mature male. Tortugas, Florida, June 21, 1907.
Figs, i to 4. LnrjJici-a cruciatn (" calcarata"). Successive stages in the

development ot' the medusa. Agassiz Laboratory, Newport,

Rhode Island. |une to September, 1892.
Fig. 5. Laodiccn cntciata ("calcarata"). Part ot" the hell-margin of the

medusa shown in figure 4.
Fig. 6. Laodicfii iriiciritii ("i nlcanita " ). Details of the gonads and manu-

bnum of the medusa shown in figure 4.
Fig. "J. Staurodiscus tctnistaunis, young medusa. Charleston Harbor, South

Carolina, September 14, 1897.
Fig. 8. Staurodiscus ti-ti-astunnis, young medusa. A sense-club from the

bell-margin.

See page I in tor description ot figure I.
Drawn from life, by the author.

PLATE 22

Pf-frf?

I 3.-

f

.>"/•/

I.LTTuMKIHs F. LMHHiT.A. 2(11

Genus LAODICEA Lesson, 1843.

l.,ia,li,ea, Lissov, 1X4;, Hist X.uoph. Aral.. i>. 21(4. \- VGA '•' m L. VcASSIz's, 1X1,2. Coin. Nat. Hi-t. I". S.. \.,l. 4,
p. 550. -Ac.Assi/ and \U\ IK, 1X99, Bull. Mils. Comp. /ool. at Harvard College, \,.l. ;2, p. ifu. H«K..III. 1904. Bull.
I .S. Bureau of Fisheries, vol. 24, p. 4^.

C.OSmftir.I, FoRKts, 1X48, British Naked-eve, ! Me. in- r, p. 42.

Thnuman'iai, GFC.SNBAI R, 1856, /cit. fur uissen. /mil., Bd. 8, p. 257.

/.<i/a?a, AC;ASSIZ, L., 1862, Cont. Nat. Hist. f. S., vol. 4, p. -,^i . A'.ASSI/, A., i Sl.s. North Ann r Veal., p. 122.

Airarfslit, \\'RII. IIT, iXhi, Ann. and Mag. Nat. Hist., ser. ;. \ol. 8. p. 129.

l'i,,/>, :lfll,i (hvdroid), HINC-KS, ]S6X, British Hvdroid /.ooph.. p. 201). plate 40.

Oelonema+ LaoJice, H^KKM, iS^, Svst. der Medu en. pp. 121"), 1^1.

/.uoilicr, MfTscHMKoth, iS8d, Embrvul. Studien an Medus.-n, \\ien, p. 23, etc. (development). -BROOKS, 1895, Journal
Morphology, vol. 10, p. 287.— BKOWM, 1896, Proc. Zool. Soc. London, p. 4X2; IhiJ.. iX.jX. p. *2;. \K*s. 1904,
Result. Camp. Sci. Prince de Monaco, Fare. z8, p. 18; 1905, Dii c ,, r> loten Meduaen der 5/laj/i Kxpedition, Monoj;. 10,
p. 24.— BF.DCIT, 1901, Revue Slliutt d.- /o,,l., tome (). p. +S;; /I,,,/., i,,o;, tome i;, p. i ;(. (,-ii.ni,,n ,,f all n
1X^0). -BKCUVNJ, l'io^. Annals and Mag. Nat. Hist., vul. 20. str. J, p. 4s"'|.

(lenu< \, MAAS, |X<(;, Hvdromedusen Plankton Kxpedition, p. ^5.

(t< >fjr fift,!//,'!, \ os. LENDENPELD, i XX 4, Proc. Linnean So. . \. • South NV.de-, vol . <f. p. ui<).

This genus was founded by Lesson, 184.5. for Lnndicfn fmiiiilii. of the coast of F.urope.
which appears to have been previously described by Forskal, 1775. under the name of M , Jam

GENERIC CM. -\RACTHRS.

Thaumandadae with 4 gonads upon the 4 radial-canals. With 4 or more tentacles. The
tentacle-bulbs often bear ectodermal ocelli. Sensory-clubs Icordylh anil cirri are situated
between the tentacles. The stomach is without a peduncle. There are 4 simple, cruciform
lips. The hvdroid is Cuspid ella Hincks.

Brooks, 180,5, shows that the sensory-clubs of /.<;«,//,,,/ are situated upon the ectodeimal
nerve-ring, which is found on the exumbrella side of the bell immediately abo\e the velum,
i he core ot each sense-club is composed ot entodermal cells which are in direct connection
with the entoderm ot the circular canal. The sense-clubs contain no concretions. Anatom-
ically the sense-clubs of Laodicea are strikingly similar to the embryonic sense-clubs of the
Narcomedusae and Trachy medusae, hut they lack concietioiis. On the other hand there can
be no doubt that Landicea is closely related to the lithoc\ st-bearing Leptomedusz or Vesic-
ulatx of the Hertwig brothers.

Haeckel introduced the spelling LaoJice, but Lesson's original name is spelled l.<i<nli, , <i.
Our Laodicea is equivalent to I.ooJici -4 Octonema Haeckel. The medusae of ].<ioJn,-n are
extremely variable in color and in the arrangement of their tentacles, cirri, ocelli, and marginal
sense-clubs. I am inclined to believe that /,. nlnt/iri\, /.. ,,il,iinitii of the Atlantic. /.. i>iJi,,i
of Ceylon, and /.. innrntnn ot the Fip Islands are only local races ot /.. crnciiitn ol the Medi-
terranean and Atlantic coasts of Europe. All of these so-called species are possibly only im-
perfectly differentiated local races of I.<i'>dic?n ,iiicnitii. Browne, iSi/i, iSc^S, has shown that
the common Mediterranean, European, and American Laodicea are probably one and tin-
same species, and my own studies serve only to convince me of the correctness of bis view.

As in T mri>f>sis the species ot Laodicea tall into two well-defined groups: 'I hose \\ith 410
8 tentacles, represented by /.. ,-;/, o/v, f,-rlilit. nrptuiin. and ^•liitinn^i. and those \\ith ^2. to 300
tentacles, represented by /,. friitintu, chapmani, and pnLhrn.

Metschmkoft, 1886, made elaborate studies ot the development of the egg of the J.nnJi. -, -it
criictntn of the Mediterranean; and he succeeded in rearing the lar\;e until tlu-\ developed into
small hydroids of the genus Cuspidella Ilincks.

Laodicea cruciata L. Agassiz.

Plate 21, figs. 4 and V: plate 22, tit;*. 2 to d; plate 2;, f\^. I to ;.

i.iwtiiffa fiilcnrtiiti, A. Anssi/, m L. A(,ASSI/'S. i SI.2. t'ouf . Nat. Hist . r . S., \ol . 4, p. ^o. H ARC. ITT, 1904, Bull. I'. S. Bureau

of Fisheries, vol. 24, p. 4^, i fit;. -.\!A\S, 1405, t'raspedoten Me.lusen ,ler Xit>og,i Expedition, Alonoi;. 10, p. 2?.
l.tifn'il rornittti, AciASSlz, L., 1862, Con:. Nat. Hist . I . S.. vol. 4, p. ^51 (non I., i i/rnulii, Lamouroux). A<.\ssi/, A., iMi;. I'

Boston Soc. Nat. Hist., vol. 9, p. 91, fii^. 4.

Caiapanularia ilumoia, LFIDV, iX^s, Marine Iim rr. Fauna of R. I. and N. J., p. f,.
l.a\a'a rulctirata, ACIASSIZ, A., iXti^, North Amer. Acal., p. 122, figs. 184, 194.
l.tioilice calfaratti, H\M KM , 1X79, Syst. der Metlusen, p. 1^4. Bkooks, i Si)^, j.iurn.il of Morpholoi;\ , vol. 10. p. :X~. plate 17,

figs. 5-7. — BROWM, iXtj^, Trans. Liverpool Biol. S..L., \,,1. 9, p. 276; 1898, Proc. Zoo]. s(ll . London for iXt>~, (•

plate 49, fig. 4; 1000, Proc. Royal Irish Acad. Dublin, ser. ;, vol. ;, p. 720.
I.afufii r,ilf,ir,a,i, VIRRIM., 187',, Report Commiss. Fish and Fishen. tot iS-i -2. p. 729.

202

MKI>U,S.<K OF TIIK WORLD.

Helirlla calcarata (hydroid and medusa), NUTTING, 1901, Bull. U. S. Fish Commission for 1899, pp. 353, 378, figs. 56, 94.
Laodice indira, BROWNE, 1905, Pearl Oyster Fisheries Gulf of Manaar, Suppl. Report 27, Roy. Soc. London, p. 136, plate i,

fig. 5; plate 4, figs. 7-11 (Ceylon).
Laodice ulothri\, HAECKEL, 1879, Syst. der Medusen, p. 133, taf. 8, fign. 5-7.— BROOKS, 1895, Amer. Journal Morphology, vol.

10, p. 304, plate 17, 7 figs.
Laodicea ultilhriv, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, (1.49; 1904, Mem.Nat.Sci. Brooklyn

Institute Museum, vol. I, p. 14, fig. 30, plate 4.
LaoJicra mamma, AGASSIZ, A., and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard College., vol. 32, p. 162, plate 3, figs. 7, 8

(Fiji Islands).

( >) Medusa ttquorea, BASTF.R, 1759, Opuscula subseciva, n, p. 55, taf. 5, fign. 2, 3.
( ?) Medusa crutiala, FHRSKAL, 1775, Descript. anim., p. 1 10, taf. 33, fig. A.

Thaumaniiai unaulata, FORBF.S and GOODSIR, 1853, Trans. Roy. Soc. Edinburgh, vol. 20, p. 313, plate 10, fig. 7.
Thiiuiiiantias medilerranea, GEGKNBAI'R, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 237, taf. 8, fign. 1-3.
l.aodi(e cruciata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 350. — HAECKEL, 1879, Syst. der Medusen, p. 132 (a list of

bibliography). — BROWNE, 1896, Proc. Zool. Soc. London, p. 483 (a good discussion of the literature). GRAEFEE, 1884,

Arbeit. Zuol. Inst. \Vien, Bd. 5, p. 357. — METSCHNIKOFF, E., 1886, Embryologische Studien an Medusen, \Vien, pp. 23

(egg)> 37 (segmentation), 57 (formation of entoderm), 83 (polypite), taf. 4, fign. 17-31; ta'' 5- ^S- '•
( r) Laodice pulchra, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 280 (Falkland Islands).
Laodicea cruciata L. \a.= Thaumanlias medilerranea GEGENCAUR; METSCHNIKOFK, E., 1886, Arbeit. Zool. Inst. \Vien, Bil . n,

p. 239.

C.osmrlira ialinarutn, or PLESSIS, 1879, Annal. Mag. Nat. Hist., London, vol. 3, p. 385 (=£. cruciata).
Laodice salinarum, HAF.CKEL, 1879, Syst. der Medusen, p. 636 (from the salt ditches of Montpellier, Mediterranean coast of

France).

Laodice cruciata, BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 483; Ibid., 1905, tome 13, p. 136 (all papers cited to 1850).
Hebella calctirala, THORNEI.V, 1905, Report Pearl Oyster Fisheries of Ceylon, Roy. Soc. London, Part 2, p. 1 16 (hydroid from

the Gulf of Manaar, Ceylon).
Laodice calcarata, etc., BROWNE, 1907, Annals and Mag. Nat. Hist., vol. 20, ser. 7, p. 460; Cuspiae/la (hydroid), Ibid., p. 463.

It is difficult to determine the correct name of this medusa. Medusa cruciata Forskal,
'775' 's so vaguel.V described and figured that it will never he possible to determine its true
relationships, and this is also true ot Medusa crrjuorcu Baster, 1759.

FH.. 104. — Laodicea " mediterranean after Gegenbaur, in Zeit. fur wissen. Zool., Bd. 8.
Oral and side views of medusa and part of bell-margin showing tentacular spurs.

Thaumanttas mediterranea Gegenbaur, 1856, is evidently a Laodicea, and is the same
medusa which L. Agassiz, 1862, and Haeckel, 1879, call Laodice cruciata. Browne, 1896,
1898, inclines to the opinion that L. calcarata, L. ulotlin\ and Thaumantias mediterranea =
L. cruciata L. Agassiz and Haeckel. and are one and the same species. My studies of
Laodicea "calcarata," which is very abundant along the Atlantic coast of the United States
south ot Cape Cod, has convinced me that it is identical with Laodicea cruciata of Europe,
and is also distributed widely over the Pacific, where it has been called L. marama and L.
indica. The extreme variability in color and in the development of the marginal appen-
dages has caused much confusion in the naming of this most abundant and widely spread
medusa. The old name L. cruciata takes precedence over "calcarata" and should supersede
it, being, indeed, synonymous with it, but nevertheless the identity between this medusa and
"Medusa cruciata" of Forskal must remain in doubt. It seems advisable, however, to retain
an old and familiar name rather than to reinstate an unfamiliar one such as /,. undulata,
which is the specific name given to this medusa by Forbes and Goodsir, 1853.

Adult medusa (plate 22, fig. 4). — The bell is about 20 to 25 mm. in diameter, and when
fully expanded it is somewhat flatter than a hemisphere. Being very flexible, however, it
assumes all sorts of distorted forms under the influence of contraction. The gelatinous sub-
stance is not thick at the apex and becomes regularly thinner toward the margin. There are
/o to 150, or more, long tentacles, the ends of which are usually coiled in a close helix, while
the main shaft of each tentacle is quite straight and rigid. The basal bulbs of the tentacles are
large and hollow and often have abaxial spur-like projections; and there is a dark-brown or

1. ATE

Figs. I and 2. LnoJii'i-n friu'iutti var. ulothnx. Successive stages in the

development <>t the medusa, Toriugas, Florida, June, i8q~.
Fig. 3. Lnndui-n cr tic nit n var. itlntlin\. Oral \'ie\v ot gonads of a female

medusa. Nassau Harbor, Bahamas, June, 1903.
Fig. 4. Mt'liiiTtuin campanula, young medusa. Nahant, Massachusetts,

May 6, 1898.
Fig. 5. Melicertum campanula, mature male. Marblehead, Massachusetts,

|ul\- n, i goo.

l^rawn from lite, bv the author.

I.KI'TOMKIM s i; I.AOIiHT \.

203

purple ocellus situated within the ectoderm of the inner (centripetal) side of" most of the ten-
tacle-bulhs. There are a number of coiled cirri scattered between the tentacle-bulbs (plate
22, fig. 5). These are usually somewhat less numerous than the tentacles. They arise at a
short distance above the bell-margin on the exumbrella suit- of the bell. The con- of each
cirrus is made up ot highly vacuolated entoderm-cells which are continuous with the entoderm
of the circular canal.

Numbers of spindle-shaped nematocyst-capsules are found in the ectoderm at the distal
end of each cirrus. In addition to the cirri there are one or two clubs, or cordyli, between each
successive pair of tentacles. The distribution of these cordvli is \ery irregular, but they are
usually somewhat more numerous than the tentacles. Kach club is flask-shaped and united
to the bell-margin by means of a narrow neck. They arise from the side of the exumbrella at
the level of the upper nerve-ring, a very short distance above the level of the velum. The core
ot each sense-club is composed ot large, highly vacuolated, entodermal cells which are directlv
continuous with the entoderm of the circular-canal (see Brooks. lS<;5. plate 17, fig. 5!. The

Fin. 105. — Ltloilicen rrntitita (ulolhrix *, .ifti-r Bnxik-. in ["iini.il i>f Mi.rpholni;\ , Vnl. 10.
A . Srai«n nl trnt.ulr. />. Sriinm nf tini- of tin- marginal (.luh-..

clubs are not solid tor there is usually a small intercellular lumen in the axis ot the broad distal
end of the club. There are no concretions. The velum iswell developed. Thereare4straighr,
narrow radial-canals. The stomach is short and quadrangular in cross-section, and there
are 4 recurved, slightly crinkled lips. In fully grown individuals the gonadj. extend from the
sides ot the stomach down nearly, it not quite, the entire length of the radial-canals. The
medusa is very variable in color, some individuals being nearly colorless. In otheis the
entoderm ot the manubnum, gonads. and tentacle-bulbs is green or pink, and the entoder-
mal lamella ot the bell is ot a delicate green, while in otheis the entoderm is milky or dull
translucent yellow.

This medusa is common along the coast of the United States from June to early winter.
along the southern shores of New F.ngland to Cuba and the West Indies, but it has not been
taken north of Massachusetts Bay. It ranges into more northerly latitudes off the European
coast, however, tor Browne took it in Valencia Harbor on the southern coast of Ireland. It
is seen in the Mediterranean from October to December, and is found in the Fiji Islands,

JIM

MEDUSAE OF THK WORLD.

and Ceylon, in the Pacific and Indian Oceans. It is common only near coasts, not being a
creature of the open ocean. It appears to be tar more abundant off the coast of America than
in European waters. I found it only occasionally at Naples during the winter of 1907 08.

The development has been studied by MetschnikofF. The early stages are described for
the Mediterranean L. cniciata. The egg is 0.18 mm. in diameter and is laid between 4 and 5
in the afternoon from November to December. Segmentation is total and equal; and an oval,
hen's-egg-shaped, ciliated, one-layered blastula is formed. Sometimes one sees a slight tem-
porary invagination at the narrow hinder end of the blastula, but this is not the beginning of the
formation of the entoderm. The entoderm on the contrary is formed from numerous cells
which migrate individually into the large central segmentation cavity from the hinder end of
the hlastula. The resulting planula swims about for a time, but finally it attaches itself and
becomes a creeping, root-like, slightly branched hydrorhiza. from which there arise unbranched
polypites "resembling Cuspidella humilis or Cuspidella costata Hincks."

The hydrotheca is long and subcylmdncal, and the margin is provided with sharp-pointed
teeth, and the polypite may withdraw within the hydrotheca and close the opening by the
folding together of these teeth. The polypites have flaring, open, urn-shaped penstomes,
below which there is a zone of jolong, slender, filiform, nematocyst-studded tentacles, all of
about the same length.

Browne, 1907, states that Miss M. Delap, of Valencia Island, off the southern coast of
Ireland, maintained a colony of Cuspidella costata Hincks alive, and obtained from it young
medusa; which resemble the youngest stages of Laodicea calcarata found in the Ocean. It is
probable, therefore, that the hydroid described by A. Agassiz, 1865, as I.afira calcarata and
which he believed to be the nurse of LarjJicea is not such. It lacks the toothed operculum
found in Cuspidella and the margin of the hydrotheca is simple and entire. The direct
embryological evidence demonstrates that the hydroid of Laodicea is CuspiJellti Hincks.

Tabular Synopsis of the "dinettes" of Ltiotliceii cruciiitn.

L.cruciata according to

L. calcarata according to mv

L. ulothrix,

L. marama

L. Jndica

Haeckel, 1879=1*.

studies.

according to

Agassiz and

Browne.

mediterranea Gegen-

Haeckel,

Mayer.

baur, 1856.

Brooks, and

Mayer.

Shape and dia-

Flat. 20 to }o

Flat to hemispherical. 20 to 25

Flat to hemi-

Flat. 5.5

Flat. 5106

meter of bell

spherical.

Cyoung).

in mm.

20

Number of ten-

100 to 120 with ocelli and

75 to 1 50 or more. Some with, 7010 looasin

50 as in L. cal-

60 to 80 with

tacles.

basal spurs.

some without, basal spurs or

L. "calcar- carala.

large ocelli

ocelli. Ocelli very variable in

ata."

and no basal

size and number. Spurs some-

spurs.

times entirely absent.

Arrangement

Clubs and cirri more nu-

Clubs and cirri sometimes more,

As in L. "cal-

As in L. "cal-

As in L. "cal-

and charac-

merous than tentacles

sometimes less, numerous than

carata."

carata."

carata."

ter of margi-

and irregularly scattered

tentacles. Sometimes quite

nal appen-

between them.

regularly arranged, hut usually

dages.

irregularly placed.

Character of

Frilled and developed

More or less frilled. Sometimes

AsinL."cal-

Immature.

Immature(?).

gonads.

along entire lengths of

mature when only about two-

carata."

Only half as

Only half as

the 4 radial-canals so as

thirds as long as radial-canals;

long as

long as

to touch ring-canal.

sometimes developed over entire

radial-

radial-

lengths of radial-canals.

canals.

canals.

Color.

Very variable. Stomach,

Verv variable, as in L. cruciata.

Variable, but

Blue to green.

(?)

gonads, and tentacle-

rarely so

bulbs faint red, purple,

brilliant as

blue, brownish, violet,

in northern

etc.

examples.

\Vhr-Tf fniind.

Mediterranean and Atlan-

Atlantic coast of United States.

Canary

Fiji Islands,

Ceylon, Indian

tic tiiavts of Europe.

Boston Harbor to West Indies.

Islands,

South Paci-

Ocean.

Bahamas,

fic.

Tortugas.

I.KITOMIJHS 1. 1. \.iDK EA.

When set free the medusa usually has 2 long tentacles, 2 well-developed hasal bulbs, and
4 small interradial swellings upon the bell-margin. At this time tin bell is about I nun. in
height and about 0.7 mm. in diameter (plate 22, rig. 2). Its outer surface is covered with
nematocyst-cells. There are only one m two marginal clubs or cirri in the young medusse.
There are, however, well-developed ocelli within the ectoderm of the inner (centripetal) side
of each tentacle-bulb. The 4 radial tubes are straight and narrow and thegonadsare repre-
sented by 4 scarcely perceptible swellings adjacent to the sides of the manubiium. The
manubrium is a simple tube with 4 small cruciform lips.

As development proceeds the tentacles increase in number, as do also the clubs and cirri.
The bell becomes relatively flatter until it approaches a hemisphere. The gonads begin to
develop at the angles ot the manubrium and soon grow down the radial-canals until they
extend to the circular canal. The manubrium acquires 4 recurved folded lips.

A synopsis ot the so-called varieties ot /.. cruciatii such as is given on page 204, may
be ot service in illustrating that we have here only one species.

Laodicea fijiana Agassiz and Mayer.

l.auJirta fjiana, AIMSSIZ, A., ami MAYFR, 1899, Bull. Mus. C'nmp. /mil. at Harvar.l (.'"llcgc, Mil. ',2, p. 163, plate 3, figs. 9, lo.
I.aotlift fijtana (var. intiim '). MAAS, 1905, CYaspr.lotcii Mrdiibrn <lrr Sibcga hij>r>l.. Munog. 10, p. 25, taf. 2, fign. 14, 15, taf.

5, fij>n. 32-35; 1906, Revue Suine de Zool., tome 14, p. Xy. BKHWM , 1907, AnnaU ami Mag. Nat. Hist., SIT. 7, vol. 20,

p. 467; L. maasi, p. 466.

Bell 12 to 20 mm. wide, 4 to 5 mm. high. Bell-walls, although thin, quite rigid. 1 60 ten-
tacles. These are flexible, tapering, with coiled ends, and are about as long as the bell-radius.
About half to three-fourths of the tentacles have a dark-brown ectodermal ocellus upon the
inner (axial) sides of their basal bulbs. There are a few clubs on the margin between the ten-
tacles; about 12 to 15 in each quadrant in a large medusa, but only about 2. in each quadrant
in specimens 6 mm. wide. There are no cirri between the tentacles.

The stomach is square in cross-section, very flat, and without a peduncle. It is about one-
fourth as wide as the bell-diameter and the lips are widely open, their folded edges inclosing
an open square.

The gonads are developed upon the proximal parts ot the 4 radial-canals and grow out-
ward from the angles ot the stomach, so that in large medusae they may extend over two-thirds
to seven-eighths of the length of each radial-canal, nearly touching the ring-canal. They are
developed upon numerous short side branches of each of the 4 radial-canals, so that they are
complexly folded and quite wide. In the female the ova stand out prominently over the surface
of the ectoderm. The outer portions of the 4 radial-canals, which lack gonads, are straight
and narrow and of the same width as the circular vessel.

The entoderm of the stomach, tentacles, and radial-canals is opaque and milky-blue in
color.

Common on the surface in the tropical Pacific, Fip, and Malay Archipelago. It apparently
does not descend into deep water.

In large medusz the wide, flat, open stomach recalls that of /}t\; liog,-mi. Maas found a
single specimen with 5 lips and 5 radial-canals, and others had 5 or (i canals. This \aiiability
recalls the condition characteristic of Dipleurosoma.

Laodicea pulchra Browne.

l.aoAict pulclira, BROWNE, 1902, Annals ami Mag. Nat. Hist., ser. 7, vol. 9, p. 280; Ibid., vol. 20, p. 466.

Umbrella flatly curved, 25 mm. wide, 15 mm. high. About 50 tentacles without basal
spurs. No marginal cirri, but j or 4 clubs (cordyli) between every two tentacles. Kach cor-
dylus situated on a small bulb. An ocellus is usually found at the base ot every tentacle and
cordylus. The stomach is very large with 4 large perradial lobes extending nearly to the
margin of the umbrella. The mouth has 4 large lips. The gonads extend from near the
stomach to within a short distance of the ring-canal. Found at Stanley Harbor, Falkland
Islands, from November to February. Color ( ?) Briefly described, without figures. b\
Browne from specimens obtained by Vallentin. May this not be identical with
crncidtfi ?

L'IMi MKDrS.E OF TIIK WOIJJ.D.

Laodicea? neptuna Mayer.

Plate 26, figs. 1-3.

f.imlirra ntfiluna, MAYER, 1900, Bull.Mus. Comp. Zoo!, at Harvard College, vol. 37, p. 48, plate 10, figs. 50-51.— BROWNE,

1907, Annals an. I Mag. Nat. Hist., ser. 7, vol. 20, p. 469.

Immature medusa (?). — Bell little higher than a hemisphere and 2.5 mm. in diameter.
There are 8 short tentacles with large basal bulbs and 8 small, rudimentary tentacle-bulbs.
Tentacles thickly covered with nematocysts and usually carried coiled and contracted. A
single, large, black ocellus is found at the base of" each tentacle (plate 26, fig. 3). There are
numerous small nematocyst-bearing cirri but no sensory-clubs upon the bell-margin between
the tentacles. The velum is well developed. There are 4 straight, radial tubes, the upper
regions of which, adjacent to the manubrium, are occupied by the gonads. The manubnum
reaches slightly beyond the velar opening and the lips are surrounded by 4 prominent clusters
of nematocyst-cells. The color of the entoderm of the manubrium, tentacle-bulbs, and circular
and radial tubes is pearly-white. The entodermal lamella of the bell is of a delicate shade of
green. This medusa was occasionally found at the Tortugas, Florida, during July and
August, 1898. It has not been seen since that time. I can not be certain that this medusa
is a l.aodicea and share the doubts expressed by Browne, 1907, upon the subject.

Laodicea? eucope.

(Irtonemti eitrv[>?, HA ECKEL, 1879, Syst. der Medusen, p. 127. — BROWNF, 1907, Annals and Mag. Nat. Hist., vol. 20, ser. 7, p. 477.

Bell hemispherical, 8 mm. wide. 8 equal tentacles, somewhat longer than the bell-
diameter and with thick, globular, basal bulbs sharply set off from the long, delicate shafts
of the tentacles. An abaxial ocellus with a subspherical lens on each tentacle-bulb. fio to 80
marginal clubs. 30 to 40 spiral cirri as in Laodicea cruclata. 4 radial-canals. Stomach with
4 short, slightly-folded lips. 4 ribbon-like, folded gonads extending along the entire lengths
of the 4 radial-canals. Color ( ?) Honolulu, Hawaiian Islands. The presence of abaxial
ocelli may necessitate the removal of this medusa from the genus Laodicea,

Laodicea? fertilis.

Ortorliofiiilon f<-'ii(i\, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 919, plate 42, figs. 14-15. —
BKOW\F, i 907, Annals and Mag. Nat. Hist., ser. 7, vol. 20, p. 478.

Rell semi-ovate, 2.5 mm. high, 2 mm. wide. Stomach a 4-sided pyramid, widest in the
middle, "octaedral." 4 simple lips. 8 tentacles, 4 radial and 4 interradial; longest tentacles
two-thirds as long as bell-height; interradial ones shorter. Tentacle-bulbs elongate, but
narrow and conical. There are 8 adradial clubs upon the bell-margin. These are longer
than the tentacle-bulbs. The gonads are complexly folded and extend over the entire lengths
ot the 4 radial-canals and fuse one with another on the sides of the stomach. Gonads orange-
yellow, other parts colorless. Sydney Harbor, New South Wales, Australia, in September.

A more detailed description is required before we can be certain of the actual affinities
of this medusa, which appears to have been described from immature specimens ?

Laodicea? chapmani Giinther.

Laoilicrn rhafimani, GiiNTHEK, 1903, Ann. anil Mag. Nat. Hist., ser. 7, vol. 11, p. 425, plate 9, figs. 1-3. North Atlantic, ofl

tin- Irish coast.
I./i'niire (?) chapmani, MAAS, 1905, Craspedoten Medusen der Sibu^n Expedition, Monog. to, p. 25. — BROWNF, 1907, Annals and

Mir. N'at. Hist., ser. 7, vol. 20, p. 469.

Bell 17 mm. wide, 12 mm. high. 32 tentacles, only 4 of which have ocelli. These 4 are
at the bases of the 4 radial-canals. Marginal clubs ( ?) Cirri ( t) 4 fimbriated, protrusive
gonads on restricted parts of the 4 radial-canals somewhat nearer to the sides of the stomach
than to the bell-margin. A single specimen taken between 1,070 fathoms and the surface off
the Irish coast in N. lat. 52° 18', W. long. 15° 54'.

The gonads do not touch the sides of the stomach. Is this an abnormal specimen of
L. cruciata or does it belong to some genus other than Laodicea?

PLATE 24.

Fig. I. Orfhistoma tentaculata, young medusa. Agassiz Laboratory, New-
port, Rhode Island, August 18, 1896.

Fig. 2. Mclifcrtissa clavigera Haeckel, Tortugas, Florida, June 2, 1906.

Fig. 3. Mcliccrtissa clavigera Haeckel. One of the marginal sense-clubs of
the medusa shown in figure 2.

Fig. 4. LnnJiffa cruciata var. ulothrtx. Part of the bell-margin of the medusa
shown in figure 2, plate 23.

Fig. 5. Mfliccrtuin campanula, halt-grown medusa with the 4 secondary
radial-canals beginning to develop. Nahant, Massachusetts,
May 6, 1897.

Drawn from life, bv the author.

PLATE 24

\

M ,

LEPTOMEDU8.*:— MELICERT1 M. L'OT

Genus MELICERTUM Oken i sensu Ehrenberg and Agassizi.

Mrliirria, preoccupied by SCHKANK, 1805, for Rotifers.— OKEN, 1815, Lehrbuch dor Naturgesch., Bd. i, Register p. 4. |'. 1:5.
PKRON rt Lt si' n'R, 1809, Ann. du Mus. Hist. Nat. Paris, tome 14, p. 352. — LESSON, 1843, Hist.Zooph. Acal., p. 324.

Mflirerlum, OKF.N, 1835, Allgemeinc Naturgesch. fiir alle Stande, Bd. 5, p. 216. — ESCHSCHOLTZ, 1829, Syst. der Acalephrn,
p. 105.— EHRF.NBERG, 1837, Abhandl. Akad., 1835, Berlin, p. 255. — AGASSIZ, L., 1862, Cont. Nat. Hist. I'. S., vol. 4,
pp. 349, 352. — AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 96; 1865, North Ainer. Acal.. p. 130. —
MAAS, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 19.-- I.I\KO, 1904,200]. Anzeigcr, Bd. 28, p. j|8.

CampiineUa, Mourn, 1857, Beskriv af Gronland, p. 95.

Melicertella -t- Melirertum, HAF.CKEL, 1879, Syst. der Medusen, pp. 134. 136.

Meliccrtidititn, BROWNK, 1901;, Proc. Royal Soc. Edinburgh, vol. 25, p. 764.

The name Melicerta appears, inadequately defined, in Oken's Lehrbuch, 1X15, on page
iv ot the index and page 125 ot the text. The same medusa is apparently described by Oken.
1X35, under the name ot Melicertum campanula from Greenland. In Oken's Melicertum
there are but 4 radial-canalsf ?) and the stomach is tubular and quadratic in cross-section
with 4 short lips. The marginal tentacles are of unequal length. It seems impossible to
determine whether the medusa described by Oken is or is not the young ot the M. campanula
of Agassi/,. The name "Melicerta" is preoccupied, and "Melicertum" would have been
hopelessly lost through inadequate definition had it not been clearly defined by A. and L.
Agassiz, 1862, 1865.

(JKNERIC CHARACTERS.

Thaumantiadae with 8 gonads upon the 8 radial-canals. 8 or more tentacles. Without
marginal sense-clubs or cirri. With or without a gastric peduncle. The hydroid is .I/,/;, ,-rtnm.

Melicertum as here defined is equivalent to M clici-ituni \ Mrli, , -it, II, i of Haeckel,
Hartlaub, 1894, and Browne, 11)05, have discovered that Oicniim 'n In, >n tutu Sars, 1835,
lacks marginal cirri and has all of the characters of Agassi/.'s Mrlicfrtum. Haeckel labored
under the false impression that this medusa had marginal clubs and he therefore established
his new genus "MelicertiJium" especially for this species. Browne would therefore amend
M elicertidium and define it in the sense of the older genus Melicertum, doing away with Melt-
cerium. Bur it appears that Ehrenberg, 1837, placed Sars's species in the genus M<-ln<-rtu>n,
and I think it should remain there and be considered a cotype of that genus.

Melicertum campanula Oken(?i, Agassiz.

Plate 23, figs. 4, 5; plate 24 fig. 5.

(?)Mrliccrtum ramfanula, OKEN, 1835, Allgemeinr Naturg. fiir Alle Stande, Bd. c, p. 226.

Mrliccrlum campanula, AGASSIZ, L., 1862, Cont. Nat. Hist. I'. S., vol. 4, pp. 349, 352 AI.ASM/,, A., |S(>;, I'm. . Huston Sen
Nat. Hist., vol. 9, p. 96, figs. 1 8, 19; 1865, North Amer. Aca!., p. 130, figs. 202-214.— HAM KM., 1879, Svst. drr Mr, I
p. 137. — Nt TTINI., 1901, Bull. U. S. Fish Commission for 1899, p. 382, fig. 104.— HARI.IT r 1904. Bull. I . S. Bureau of
1'isherirs, vol. 24, p. 44. — I.INKO, 1904, Zool. An/eiger, Bd. 28, p. 2|8.

Mftitertidium {(ittipaiiula, BROWNK, 1905, Proc. Royal Soc. Edinburgh, vol. 2s", p. 766.

Adult infiiusa (plate 23, fig. 5). — Bell pyntorm in outline and about 25 mm. in height and
25 mm. in diameter. The gelatinous substance is quite thick, especially at the aboral pole.
There are about 70 long, flexible tentacles with hollow, conical, basal bulbs. The tentacles
arise from the hell-margin and are all longer than the bell-height. There are no marginal
cirri or lithocysts between the tentacles, nor any ocelli. The velum is well-developed. There
are 8 simple, straight, quite broad radial tubes. The manubnum is wide and shallow and
does not extend quite half the distance from the inner apex ot the bell-cavity to the velar open-
ing. There are 8 simple, recurved lips. The 8 gonads occupy the entire lengths of the S
radial-canals. These are longer than the canals upon which they lie and are reflected from one
side to the other of the radial-canals in sinusoidal curves. The entoderm of the manubiium,
gonads, and tentacle-bulbs is ocher-yellow, and the bell often displays a taint yellow tinge.

H \drold and \ouno medusa. — The eggs are cast into the water and develop into pear-
shaped planulae. The planulae are ciliated and swim about near the bottom of the aquarium.
They soon attach themselves by the blunt (foremost) end. and the narrow ((undermost) end
then develops into a long, narrow trunk, which becomes the mam stem of the hydroid. The
distal extremity of the stem expands and forms the first polvpite. The polypites are long.
cylindrical, and protected by a small, funnel-shaped hydrotheca at their proximal ends.

20S MKIK'S-K OF THE WORLD.

There is a circlet of 10 to 12 short, stout tentacles, the bases of which are connected by a web.
At the end of six months the hydroid colonies were about 8 mm. in height.

In the youngest medusa observed the bell is about twice as high as broad and the bell-
walls are thin and flexible. There are 2 well-developed, diametrically opposite tentacles and
2 small, undeveloped tentacle-bulbs. There are 4 straight, radial tubes. The manubrium is
very small and there are 4 simple lips.

When the medusa is 1.5 mm. in diameter (plate 23, fig. 4) the bell has become relatively
flatter and broader. There are now 8 tentacles, 4 long radial and 4 short interradial ones.
There are, as yet, but 4 radial tubes and 4 lips. The bell-walls are still thin.

When the medusa is about 4 mm. in diameter (plate 24, fig. 5), the bell has become
pvriform in shape and the gelatinous substance has greatly increased in thickness. There
are }2 tentacles in several stages of development. There are 8 radial-tubes, 4 of which are
broad, and 4 very narrow and almost thread-like. The 4 thread-like tubes are new and arise
from the periphery of the stomach and extend down toward the circular vessel. The manu-
brium is shallow and there are 8 simple lips. The gonads begin at this stage upon the 4 broad
radial-canals. The other 4 radial-canals acquire them later. The gonads first appear near the
periphery of the stomach and extend down the radial-canals until they reach the circular tube.

This medusa is common on the New England coast, north of Cape Cod, during May and
early ] une. It has not been taken south of Cape Cod. It is exceedingly abundant on the sur-
face in Salem Harbor, Massachusetts, early in the summer, but disappears about the middle
of July. The medusa is very constant to its type, individuals having more or less than 8
radial-canals being exceedingly rare. Lmko finds it to be rare off the Murman coast, Northern
Russia, near Norway.

Mehcertum oi'torostntiint ( = 0ceania octocostnta Sars) of the northern Atlantic coasts of
Europe is closely allied to, if not identical with, our American medusa.

Melicertum octocostatum Haeckel.

Oimntti tjilfjcottatti, SARS, 1831;, Beskriv og Jagttag, p. 24, plate 4, fig. 9 «-</.

\tfliif>ntm campanulalum, EHRENBERG, 1837, Abhandl. d. k. Akad. Wissen. zu Berlin (1835), pp. 190, 255, taf. 8, fign. 5-7-

Thaumantias mtlltri, LANDSBOROUGH, 1847, Arran, a poem, p. 265.

Stomobrachium octocostatum, FORBES, 1848, British Naked-eyed Medusae, p. 30, plate 4, fig. I . — ROMANES, 1876, Journal Linnean

SMC. London, vol. 12, p. 526.

Mclicfrtum octocostalurri, Mcltctrtidium oclofottatutn, HAK.CIO.L, 1879, Syst. der Medusen, p. 138.
Mi-la frlnlium octocostatum, HARTLAUB, 1894, Wissen. Meeresuntersuch. Kornm. Meere Kiel, Helgoland, Neue Folge, Bd. i, p.

192. — BROWNE, 1905 Proc. Roy. Soc. Edinburgh, vol. 25, p. 762. — BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 138

(literature 1835-50).

Bell somewhat conical to pyntorm, with thick, solid apex and thinner sides. About I I
to 13 mm. high and 10 to 1 1 mm. wide. There are about 64 large, alternating with 64 small,
tentacles having laterally compressed basal bulbs and no ocelli. No clubs or cirri. Velum
narrow. There are 3 to 5 fine, longitudinal lines of nematocyst ( :) cells upon the subumbrella,
extending from stomach to margin between each successive pair of radial-canals. These lines
occasionally branch or anastomose. In large medusae these lines are usually confined to parts
near the margin, extending upward for a short distance only. Such lines have never been
observed in the American M. campanula.

The stomach is octagonal, prismatic, one-fourth as long as the depth of the bell-cavity.
It has 8 slightly crinkled lips. A short, broad peduncle is sometimes present. The 8 radial-
canals are broad and straight, and the sinuous linear gonads extend along their outer two-
thirds, reaching the bell-margin.

Color of gonads, stomach, and basal bulbs of large tentacles, yellow to yellowish-brown.

Found ofF the northern Atlantic coasts of Europe, Norway, Helgoland, Scotland. The
best modern descriptions are given by Browne and Hartlaub.

This medusa appears to differ from the American M . campanula Agassiz in having fine
radiating lines of nematocysts over its subumbrella. The gonads appear to be more re-
stricted in their position upon the radial-canals, and the tentacles to be more markedly of two
sr/.es in the European form. Nevertheless the two "species" are evidently closely related and
may eventually prove to be identical. This is rendered probable on account of the Arctic
range of M . anufiuuula Agassiz.

LKPTOMEDUS.fi — MF.LICKiriVM, M K LICKHTISS \. •_>()()

Melicertum georgicum A. Agassiz.

Melicertum georgif urn, \CASSIZ, A.., in I,. Agassiz's, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 349; 1865, North American Ac.il,
p. 135, figs. 115, 216.— HAECKEL, 1879, Syst. der Medusen, p. 137.

Bell pointed above; 20 mm. high, 20 mm. wide. About 30 to 40 tentacles with large
basal bulbs. No peduncle. Stomach wide, flat, and octagonal. Mouth long, with 4 ( ' r) lips.
8 gonads covering nearly the entire lengths of the 8 radial-canals, but not touching the ring-
canal. Manubrium, gonads, and tentacle-bulbs yellow.

Pacific coast of North America, State of Washington, in July.

Distinguished by its small number of tentacles and 4 ( ?) lips. Is it identical with M.
campanula Agassiz ?

Melicertum panocto.
Mtlicertflla panocto, HAECKEL, 1879, Syst. der Medusen, p. 135.

Bell is high, rounded, nearly barrel-shaped, 10 mm. high, 8 mm. wide. No marginal
clubs or cirri. 8 tentacles longer than bell-diameter, with very large, globular basal bulbs.
A black ocellus on each tentacle-bulb. The ocellus is flanked by a half-moon-shaped black
spot on both sides. Stomach octagonal, prismatic, half as long as bell-height. 8 long, much
folded, crinkled lips. 8 spindle-shaped gonads developed upon entire lengths of 8 radial-
canals. Color ( ?) Azores, Atlantic.

Described by Haeckel from a preserved specimen. May it not be a specimen of .\f,li-
certissa clavigera which has lost its marginal clubs ?

Melicertum (?) Aglantha (?) proboscifer Maas.
Mcliccrtum proboscifer, MAAS, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 13, p. 19, taf. z, fign. 5-7.

Bell rounded, 40 mm. wide, 15 mm. high; bell-walls quite thin. No apical projection.
About 1 60 or more short, slender, tapering tentacles, nearly all of which were broken off
short in the specimen studied by Maas. No lithocysts observed. Velum weaklv developed.
8 straight, narrow radial-canals and a simple, marginal ring-canal. Stomach small, urn-
shaped, and mounted upon a narrow, long, cylindrical peduncle as long as depth of bell-
cavity so that the stomach projects beyond velar opening. There are 8 short, recurved,
somewhat folded lips. 8 sausage-shaped gonads arise from middle of subumbrella lengths of
the 8 radial-canals and project outward into bell-cavity.

Entodermal parts, including interstitial lamella of bell, intense carmine; tentacles
rose-colored.

Found in the Gulf of Panama, Pacific coast of Central America, in March, 1891, by the
Albatross.

I believe that future studies will show that this medusa is one of the Aglauridae, possi-
bly Aglantha. Its slender (solid ?), brittle tentacles, 8 sausage-shaped gonads, and lon<;,
cylindrical peduncle are all characters of Aglant/ia. The lithocysts of Aglauridae are often dif-
ficult to observe owing to their being readily lost, even in living medusae, and in preserved
specimens they are rarely detected. All known species of A yluntha have 4 lips, but this medusa
is said to have 8; however, its general appearance is wholly unlike that of any other Meli-
certum, but closely resembles that of the known species of Aglantlia.

Genus MELICERTISSA Haeckel, 1879, sens, emend.

\lelicertissa+ Melicrrlitlium, HAECKEL, 1879, Syst. tier MeJusen, pp. 124, i;c, 137.
Mflicerltjiurn, MAAS, 1905, CraspcdotrM MrdiiM-n der Siboga Expedition, Monog. lo, p. IJ.
Melicertiisa, BROWNF, 1907, Annals and Mag. Nat. Hist., ser. 7, vol. zo, p. 476.

CKNKRIC CHARACTKRS.

Thaumantiadae with S simple radial-canals upon which the 8 gonads are developed.
8 lips and 8 or more marginal tentacles. Marginal sense-clubs or cirri, or both, are present.
Development unknown.

'7

210

MEDUS.fi OF THE WORLD.

Our Melicertissa is equivalent to Melicertissa -\-Melicertidium of Haeckel, 1879. Haeckel
would restrict Melicertissa to include medusae with only 8 tentacles, while Melicertidium would
apply to medusx with more than 8 tentacles. It is probable that Melicertissa as defined by
Haeckel is only the young of his Mi-liiertiJiiim. His name Melicertissa precedes that of Mel i-
certiJiuin and should therefore designate the genus.

Melicertissa is closely related to Melici-rttitn, but is distinguished by its rudimentary mar-
ginal sense-clubs or cirri which are absent in Melicertutn.

Melicertissa clavigera Haeckel.
Plate 24, figs. 2 and 3.

Melictriiisa clavigera, HAECKEL, 1879, Syst. der Medusen, p. 135, taf. 8, figs. 8-12.

( ?) Melicertella panoclo, HAECKEL, Ibid. p. 135 (a damaged specimen without marginal clubs?).

Bell 7 mm. in diameter, flatter than a hemisphere, with moderately thin walls becoming
thinner toward margin. 16 marginal tentacles with long, hollow, tapering basal bulbs and
coiled lash-like ends. When contracted these tentacles are each about half as long as the bell-
diameter. 8 of the tentacles are radial and 8 interradial. There are 16 sense-clubs (plate 24,
fig. 3) midway between the tentacles. There is a well-developed, black, entodermal, cup-like
mass of pigment at the base of each tentacle and sense-club, 32 in all. In some specimens
one or more of the interradial tentacles are replaced by a marginal sense-club. Haeckel, indeed,
describes only 8 radial tentacles and 24 marginal sense-clubs, there being no interradial ten-
tacles. His specimens were obtained in the Canary Islands. At Tortugas, Florida, however,
we find specimens with interradial tentacles, but which conform in all other respects toHaeckel's
medusa.

106.

FIG. 106. — Melicertissa clavigera, after Haeckel, 1879.

FII;. 107. — Melicertissa malayica, after Maas, in Hydromedusen Siboga Expedition.

Velum well developed. There are 8 straight, narrow radial-canals and a simple, circular
vessel. Stomach about one-fourth as wide as bell-diameter and the 8 radial-canals arise singly
from its periphery 45° apart. Stomach flat and entire manubrium short and not over one-
third as long as depth of bell-cavity. There are 8 short, simple lips. The gonads are linear,
somewhat sinuous, and occupy the middle halves of the radial-canals, not touching the cir-
cular canal or edge of stomach.

Entoderm of stomach, radial and circular canals, and tentacle-bulbs dull yellow in speci-
mens from Tortugas, Florida ; the entodermal marginal ocelli black. Haeckel describes the
color of the entoderm in the Canary Island specimens as being bronze-colored, brownish, or
greenish-yellow.

Haeckel found this medusa at Lanzerote, Canary Islands, in January, 1867. We found it
on the surface at Tortugas, Florida, early in June, 1906.

The American specimens are interesting, for they show that there is no natural distinc-
tion between the genera Melicertissa and Melicertidium of Haeckel. Parts of the bell-rim are

LEPTOMEDUS.E— MKLICEUTISSA, ORfHISTOMA. 211

usually in the " Melicertissa" condition with only 2 tentacles and 3 sense-clubs in each octant,
while other parts of the rim of the same medusa are in the " Melicertidium " condition with 3
tentacles and 2 sense-clubs in each octant.

Melicertissa malayica Browne.

Melicertidium malayicum, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 28, taf. 5, fign. 19-31.
Melicertissa malayicat BROWNE, 1907, Annals and Mag. Nat. Hist., scr. 7, vol. 20, p. 477.

Bell flat, 32 mm. wide, 20 short tentacles in each octant, 160 in all. These tentacles are
only about one-tenth as long as the bell-radius and their outer ends are coiled. They have
slightly swollen basal bulbs. At the base of about every fourth tentacle there is an abaxial,
ectodermal, swollen ocellus. Altogether there are 40 to 48 of these ocelli.

A number of small clubs arise from the margin between the tentacles. These are about
hall as numerous as the tentacles and are irregularly scattered around the margin. The cores
of these clubs are filled with entodermal cells, as in LaoJicea. A few small, coiled cirri, much
fewer than the clubs, arise from the margin, each containing an axial core of entoderm.

Velum narrow. There are 8 straight, narrow radial-canals and a simple, slender ring-
canal. Stomach flat, without a peduncle and with 8 simple, folded lips. Gonads linear and
developed upon the sides of the proximal thirds of the 8 radial-canals, adjacent to stomach.
The gonads begin to develop when the medusa is 18 mm. wide. The ocelli at the bases of the
tentacles are black. (See figure 107, page 210.)

Malay Archipelago, Ternate. 2 specimens found by the Siboga expedition.

Genus ORCHISTOMA Haeckel, 1879.
Orchisioma, HAECKKL, 1879, Syst. der Medusen, p. 138.— KELLER, 1884, Recueil Zool. Suissc, tome I, pp. 417, 418.

The type species of this genus is Orchistorna pileus, described by Lesson (1845, p. 317,
plate 6, fig. i) under the name of Mesonema piletis. This species is believed to have come
from the west coast of Africa.

GENERIC CHARACTERS.

Thaumantiadz with more than 8 simple radial-canals which arise separately from periph-
ery of stomach. The gonads are situated upon the radial-canals. Numerous tentacles.
Cirri or marginal sense-clubs may or may not be present.

Mesonema of Eschscholtz, 1829, is one of the yEquoridae, and hence this generic name can
not be applied to the type species of Orchistorna.

Orchistoma pileus Lesson.

Plate 25, figs, l to 4.

Mesonema pileus, LESSON, 1843, Hist. Zooph. Acal., p. 317, planchr 6, fig. i.

Orchisioma steenstrupii, HAEI KEL, 1879, Syst. der Medusen, p. 139, taf. 15, fign. 3-5.— KELLER, 1884, Recueil Zool. Suisse, tome
I, p. 417.

Bell flat-topped with vertical sides, 30 to 40 mm. in diameter. Gelatinous substance
thick. 64 short tentacles with well-developed, hollow, basal bulbs; these tentacles are shorter
than the bell-radius and are flat and ribbon-like, with hollow bulbs. About 100 small, solid,
straight cirri from the side of exumbrella at a short distance above the bell-margin (plate 25,
fig. 2). The entodermal cores of these cirri are connected with the entoderm of tin- circular
canal. There are about 400 dark-brown, entodermal ocelli upon the circular canal. Each
ocellus is provided with an ectodermal lens (plate 25, fig. 3).

Velum narrow. There are 32 straight, narrow radial-canals. The manubrium is pro-
vided with a large peduncle fully half as wide as bell-diameter and not quite as long as wide;
it extends a short distance beyond the velar opening. The stomach is wide and shallow, and
there are 32 long, complexly crenated lips (plate 25, fig. 4). There are 32 short, swollen,
linear gonads, one upon each radial-canal very near its point of juncture with the stomach.

In the female the gonads are cinnamon-color, lips sage-green, tentacles ocher-yellow, and
ocelli black. Other parts colorless. In the male, however, gonads, lips, and tentacle-bulbs
are dull blue-gray.

212 MEDUS.E OF THE WORLD.

Lesson first found this medusa off the west coast of Africa ( ?). Haeckel describes it from
the West Indies, and I have taken it upon the surface of the Gulf Stream off the Bahamas and
near Tortugas, Florida, from March to July. The males greatly outnumber the females.

Distinguished from O. agariciforme Keller, from the Mediterranean, by having 32 lips
instead of only 7, as in O. agariciforme.

Usually very rare, but abundant during the spring and summer of 1907, at Tortugas,
Florida.

Orchistoma agariciforme Keller.
Orchistoma agariciforme, KELLER, 1884, Recueil Zool. Suisse, tome I, p. 418, taf. zi, fign. 1-3.

Found by Keller at Naples, Italy. Similar to Orchistoma pileus in all respects except
that there are but 7 lips instead of 32, as in the Atlantic species. The color is not described
by Keller, who found only one mature and one young specimen. Like the Atlantic form, it is
very rare. 19 or more (about 32 ?) radial-canals. Keller states that the gonads are diverticula
from the stomach which extend outward along the radial-canals.

Orchistoma tentaculata Mayer.

Plate 24. fig. i.

Orchntoma tentaculata, MAYER, 1900, Bull. Mus. Comp.Zool. at Harvard College, vol. 37, p. 8, plate 5, fig. 19. — NUTTING, 1901 ,
Bull. U. S. Fish Com. for 1899, p. 377, fig. 92. — HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 44, I fig.

Youn% medusa. — Bell 6 mm. in height. Sides near margin slightly flanged outward.
Gelatinous substance of upper part of bell very thick, so that concavity is shallow. 32 mar-
ginal tentacles in various stages of development, the longest being about 1.5 times as long
as bell-height. Tentacles with long, hollow basal bulbs. No marginal sense-organs. 16
functional radial tubes and 16 others in process of development. Radial tubes straight with
no traces of gonads upon them. Velum well developed. Manubrium flat and shallow, 8 lips.

The entoderm of the basal bulbs of the tentacles delicate green.

Only one immature specimen of this medusa has ever been seen; no gonads were visible.
It was found on the surface at Newport, Rhode Island, August 18, 1896.

Nutting states that the tentacle-bulbs are red. They are certainly green by reflected
light, but probably red by transmitted light. Descriptions of color in this work are based
upon the appearance presented by reflected light. Professor Nutting's figure and mine are
drawn independently from the same specimen while yet alive.

The differences between this medusa and 0. pileus are so considerable that it does not
seem possible that the one can be the young of the other.

Genus TIMOIDES Bigelow, 1904.

Timoides, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 254.

The only known species is Timoides agassizn Bigelow, of the Maldive Islands, Indian
Ocean.

GENERIC CHARACTERS.

Thaumantiad.-c with 4 radial-canals and a ring-canal which gives rise to blindly ending,
centripetal canals. Stomach mounted upon a peduncle. Tentacles numerous, without lateral
cirri. Numerous cirri upon bell-margin. There are no lithocysts. 4 gonads on the 4 radial-
canals. Dr. Bigelow, upon re-examining his 'specimens, tells me that the medusa has no
lithocysts.

Timoides agassizii Bigelow.
TimoiJes agassizii, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 254, plate 3, figs. 10, n.

Medusa bell-shaped with rounded dome and flaring sides; 20 mm. wide, 14 mm. high,
with very thick gelatinous substance at apex, but thin at margin. 32 tentacles, 4 radial, 4
interradial, 8 adradial, and 16 intermediate. The 4 radial tentacles are at the ends of the 4
radial-canals. The 4 interradial and 8 adradial tentacles are at the bases of the 12 interradial

LKI'TOMKIH'S.K TlMnlDKS, STAl KODIsrlS.

21:5

and adradial centripetal canals which arise from the ring-canal and end Mindly in the gelat-
inous substance of the bell. The remaining 16 tentacles alternate with the radial and
centripetal canals. All of these tentacles are exceedingly flexible and contractile; when
contracted they are usually coiled helically, this coiling being more or less local and usually
not affecting the entire length of the tentacle. When expanded, the tentacles are about 300
mm. long. In addition to the tentacles there are numerous cirri along the bell-margin, but no
cirri arise from the sides of the tentacle-bulbs. No lithocysts upon the bell-margin.

The 4 radial-canals are straight and slender. There is a marginal ring-canal which gives
off 12 blindly ending, centripetal vessels, the 4 interradial ones being about two-thirds as long
as the bell-height, and the 8 adradial ones are only half this length. Velum well developed.

Peduncle nearly cylindrical, flaring at its base, and
twice as long as depth of bell-cavity. The barrel-shaped
stomach is almost as long as the peduncle and bears
4 lancet-shaped lips with complexly folded margins.
These lips may be longer than the combined lengths of
stomach and peduncle.

The 4 gonads are upon the lower half of the
peduncle on the 4 radial-canals, above the stomach.
I hey consist of a great number of simple and branched
papilliform processes on both sides of the 4 radial-
canals, thus forming 4 prominent, double ridges.

The gelatinous substance of the hell is faintlv tinged
with blue. The radial-canals and tentacles are rose-
pink and there is a pink pigment-spot at base of each
tentacle. Stomach and lips pink-\iolet. Gonads rich
Indian yellow, changing in some lights to ruddy-orange.
I his beautiful medusa was found by Dr. Bigelow
in abundance in Haddummati Atoll, Maldive Islands.
Indian Ocean, on January 8, 1902. It appears upon
the surface at about 4 p. m.

Genus STAURODISCUS Haeckel, 1879.

Staurodisfus, HAECKEL, 1879, Syst. der Meilusrn, p. 145. — BROWNE, 1907,
Annals and Mag. Nat. Hist., SIT. 7, vol. 20, p. 475.

\on Staurodixus, AGASSIZ AND MAYJR, 1X99, Bull. Mus. Comp. Zool. at
Harvard College, vol. 32, p. 164.

Staurodiscua (in part), MAAS, 1904, Sit/unr her . ni.itli.-pli\ > . Kl.issr ki;l.
Bayer. Akad. der Wissenschaflcn, Bd. ^4, p. 440.

This genus was founded by Haeckel, 1874, for
StnurrtJisfU) t.-timtiiurus found at the Canary Islands
and off the Honda coast.

FK;. 108. — Timoides agassizit, after BIL'''"'A,
in Bull. Museum Comp. Zoo], at
Harvard College.

(.1 NFRIC CHAKACT1 KS.

Thaumantiadae with 4 radial-canals, each Diving
rise to 2 side branches which end blindly. Tin- uonads
are upon the 4 principal radial-canals and their side branches. There may or may not be
marginal sense-clubs between the tentacles. Development unknown.

Staurodiscus is closely related to Cannot a, but in duniiotn the side branches of the radial-
canals fuse with the circular vessel, whereas in Staurodiscus they end blindly. Maas, 1404,
considers Staurodiscus to be the young of ('./niiinin. and believes that the side branches
of the radial-canals eventually fuse with the ring-canal. I have seen hundreds of medus;e of
Staurodiscus ti-tniituurus, evidently mature, but have never found the side branches fused with
the ring-canal, ('.unnotn doubtless passes through a "Staurodiscus stage," but on the other
hand there appear to be medusae which do not attain to the C.nnnntii condition anil must be
called Staurodiscus. "Staurodiscus" mgrifans Agassi/ and Mayer should be called C

214 MEDUSAE OF THE WORLD.

nii;ri<:ans. It is probable that Dipleurosorna brooksn is derived from a Staurodiscus
ancestor by tbe new formation of 4 interradial canals and the growing outward of the side
branches of the perradial canals until they join the ring-canal.

Staurodiscus tetrastaurus Haeckel.
Plate 22, figs. 7 and 8; plate 25, fig. 5; plate 26, figs. 10 and 11.

SlaaroJiscui tetraslaurus, HAECKF.L, 1879, Syst. der Medusen, p. 145, taf. 9, fign. 1-3. — MAYER, 1900, Bull. Mus. Comp. Zool.

at Harvard College, vol. 37, p. 46, plates 1 8, 19, figs. 47-49. — MAAS, 1904, Sitzungsber. math . -phys. Klasse kg). Bayer. Akad.

der Wissenschaft., Bd. 34, p. 440. — BROWNE, 1907, Annals and Mag. Nat. Hist., ser. 7, vol. 20, p. 475.
Ottarrftna gelalinosa (young medusa), MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 8, plate 6, figs.

20, 2 1 .

Bell 4.5 mm. in diameter, and about twice as wide as high. In adult medusae there are 8
long, flexible tentacles with hollow basal bulbs. Some specimens have 24, others 16 sensory-
clubs upon the bell-margin. 32 black entodermal ocelli, one at the base of each tentacle-bulb
and sensory-club. Velum is well developed. Only 4 radial-canals reach the circular vessel.
From the middle points of each of these canals there arises a pair of equally developed side
branches that end blindly. The gonads are upon these side branches and on the distal
part of each radial-canal. The manubrium bears 4 prominent lips. Entoderm green or yellow.

In the youngest specimen observed, the bell was I mm. in diameter and about as high as
broad. There were 4 well-developed tentacles, 4 rudimentary tentacle-bulbs, and 8 marginal
clubs. The medusa is common on the surface at Tortugas, Florida, in July and August.
Haeckel found this species in the Canary Islands, at Lanzerote. I have seen hundreds of
specimens at Tortugas and the side branches of the 4 radial-canals appear never to extend
to the circular vessel.

Staurodiscus heterosceles Haeckel.

Staurodiscus heterosceles, HAF.CKEL, 1879, Syst. der Medusen, p. 146.

This medusa resembles S. tetrastaurus Haeckel, excepting that the two side branches from
each of the 4 radial-canals do not arise opposite one another from the middle of the radial-
canals, but one branch is long and arises from the main canal near the stomach, while the other
side branch is short and arises from the main canal beyond the middle point near the bell-
margin. The long branches arise constantly from one, and the short branches from the
opposite sides of the main radial-canals as one glances around the bell.

Bell hemispherical, 6 to 8 mm. wide. There are 8 to 3*, usually 1 6, long tentacles with
globular basal bulbs, and 40 to 80 marginal clubs between the tentacles. There is an ocellus
at the base of each tentacle-bulb and marginal club. 12 club-shaped gonads, one at the
outer end of each of the 4 main radial-canals, and one upon each side branch. Stomach,
gonads, and tentacle-bulbs yellowish-brown. Ocelli black. Canary Islands, January, 1867,
Haeckel. Development unknown.

Genus PTYCHOGENA A. Agassiz, 1865.

I'lwhogena, Ar.ASSiz, A., 1865, North Amer. Acal., p. 137. — HAECKEL, 1879, Syst. der Medusen, p. 147; 1881, Report on Deep-
sea Medusa?, Challenger Report, Zool., vol. 4, part xn, p. 7.

\ r»i Ptychogena, MAAS, 1893, Ergeb. der Plankton Exped., Bd. u. K. c., p. 64.

I'lytliogetia, LEVINSEN, 1893, Vid. Meddel. Nat. For. Kjbbenhavn, ser. 5, Bd. 4, p. 145. — LINKO, A., 1904, Zool. Anzeiger, Bd.
28, p. 217. — BROWNE, 1907, Annals and Mag. Nat. Hist. ser. 7, vol. 20, p. 473. — BICELOW, 1909, Mem. Mus. Comp.
Zool. at Harvard College, vol. 37, p. 149. — TORREY, 1909, University California Publications, Zool., vol. 6, p. 13.

This genus was founded by A. Agassiz, 1865, for Ptychogena lactea, found in Massachu-
setts Bay.

GENERIC CHARACTERS.

Thaumantiadae with 4 radial-canals which in the mature medusa give rise to numerous,
short, blindly ending side branches. The gonads are situated upon the radial-canals in the
interspaces between the side branches, close to the suhumbrella. The stomach is a wide,
Hat pouch and the mouth is a simple cruciform slit. The tentacles have no ocelli, and differ
in this respect from those of I. nod icca.

PLATE 25.

Fig. i .Orchistoma pileus, mature medusa, female. Gult Stream, Tortugas,

Florida, July 4, 1906.

Fig. 2. Orchistoma pileus. Part of the bell-margin.
Fig. 3. Orchistoma pileus. One of the ocelli showing the ectodermal doubly

convex lens and the entodermal pigment.
Fig. 4. Orchistoma pileus. One of the lips.
Fig. 5. StdiirrjJiscus tetrastaurus. Side view of mature male. Tortugas,

Florida, August, 1898.
Fig. 6. Staurophora mertensii, young medusa. Nahant, Massachusetts,

May 7, 1897.
Figs. 7 and 8. Dipleurosoma collapsa, young stage in the growth of the

medusa. Tortugas, Florida, and Nassau, Bahamas, July, 1902-03.

Drawn from life, by the author.

PLATE 25

LEPTOMKDUS/E — PTYCIIU< MA \. - 1 .">

Ptychogena lactea A. Agassiz.

Ptychogena lac tea, AGASSIZ, A., 1865, North Amor. Acal., p. 137, figs. 220-224; 1888, Bull. Mus. Comp.Zool.at Harvard College,
vol. 15, p. 128, fig. 422. — HAECKEI., 1879, Syst JIT Mcdusen, p. 147.— BROWNE, 1907, Annals anJ Mag. N.it. Hi ' .MT. 7,
vol. 20, p. 473.

Ptychogena fiinnulata, HASCKF.L, 1879, Syst. Jcr Meduscn, p. 148; 1881, Rrport on Dccp-sea Medusa-, Report^ H. M. S. Chal-
lenger Zool., vol. 4, part ] 2, p. 7, plate 2, figs. 1 lo 8. — LEVINSEN, 1893, Vid.Meddel. Nat. Fur. Kjcbtnhavn, ser. 5, Bd. 4,
p. 145. — GRUNBLRG, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 11, p. 465.

Plychogrna finnulata var. intermedia LINKO, 1904, Zool. Anzeiger, Bd. 28, p. 217.

Adult medusa. — Bell about 50 to 70 mm. in diameter and 20 to 30 mm. in height. Sides
ot bell flange slightly outward. The gelatinous substance is about 10 mm. thick at aboral pole
of bell, but diminishes constantly in thickness towards the margin. Substance of" bell trans-
parent, but quite rigid. There are more than 300 long, slender, highly contractile tentacles,
with hollow, finger-shaped basal bulbs. When expanded these tentacles are about as long
as the bell-diameter. Their free extremities often become entangled and this causes the
tentacles to droop in graceful festoons from the hell-margin. There are 25 to 200 or more
marginal sense-clubs. They arise on the inner (centripetal) side of the tentacle bases near
the origin of the velum. These sense-clubs are elongate and pynform, with a narrow basal
stalk. They are between a third and a half as long as the basal bulbs of the tentacles.
Each sense-club is hollow and connected with the lumen of the circular canal. In some
places the sense-clubs are as numerous as the tentacles and often alternate with them in
position, but on other parts of the bell-margin they are much less numerous than the tentacles.
Velum very wide and thin. The 4 radial-canals are wide near their proximal ends, ad|acent to
manubrium, but straight and narrow near the circular vessel. These radial-canals are pin-
nated in their proximal halves, adjacent to the stomach. In this region they give rise to about
20 to 30 blindly ending, pinnate side branches, upon which the sjenital organs are situated.
The genital organs consist of leal-shaped sacs attached to the side branches of the radial-
canals. The free outer edges of these sacs are serrated or split up into a number of finger-
shaped points. Thus each genital gland consists of 20 to 30 leaf-shaped sacs or lamellae, all
connected with the main axis of the radial-canal. Manubrium wide and shallow. The 4 corners
of the fundus of the stomach are prolonged into 4 conical tunnels, whose ends extend to the
middle of the genital organs and occupy the proximal halves of the 4 radial-canals. Mouth-
opening quadrate and about one-third as wide as bell-diameter. Lips irregularly trilled and
extend at the 4 radial corners into 4 short, wavy, oral lobes. 4 narrow, ciliated grooves extend
along umbrella wall of the 4 radial-canals and meet at center of stomach on gastral surface of
gelatinous umbrella, forming a cross. The gonads and tentacle-bulbs are milk)' in color and
the marginal sense-clubs are white. All other parts of the medusa are transparent.

This medusa appears to be an Arctic form. It was first found by A. Agassiz swimming at
considerable depth in Massachusetts Bay. Since then specimens have been described by
Haeckel, under the name of Pt\clws>t-ii<i pinuuliita, from the North Atlantic Ocean between
Ireland and Iceland (lat. 50° f N., long. 13° 32' W. from Greenwich). The Challenger also
dredged a specimen between 1,250 fathoms and the surface near Halifax, Nova Scotia. Lev-
insen, 1893, records this medusa from Jakobshavn and Ritenbenk on the west coast of
Greenland, and Linko found it in Barents Sea in 72° 30' N. lat., both in the eastern and west-
ern parts of the sea north ot Lapland.

I believe that P. fiiiniilntii of Haeckel, 1879, is identical with P. luitea of A. Agassiz, 1865.
Upon comparing Agassiz's original figures (some of them unpublished) with those of Haeckel,
it appears that the only difference which may exist between the two forms is that in /'. piiinu-
l/itii the free, outer edges of the leaf-like gonads are serrated, whereas in P. Inctea of Agassiz
they appear to be entire and smooth. Also in Agassiz's figures some of the side branches of
the radial-canals give rise to secondary branches, and this appears not to be the case in
Haeckel's /•*. fiiniulntn. Agassi/.'s specimens were, however, larger than those described
by Haeckel, and probably they may have been more matuu-.

Linko, 1904, found a medusa in Barents Sea which he considers to be intermediate
between P. Itiftfd and P. pinnulutu, but nearer to I', finnulata.

It is possible that this medusa is an inhabitant ot the deep-sea and that it comes to the
surface only occasionally.

216

MEDUSAE OP THE WORLD.

As A. Agassiz says:

"The action of the light and increase of temperature of the surface is sufficient to kill them in
the course of half an hour; the moment they are brought to the surface, the bell loses its transpar-
ency, the genital organs become dull and the medusa is soon completely decomposed. This action
is much more rapid than anything of the kind which I have noticed, even in Ctenophora;, Mertensin
being the only genus in which the decomposing effects of light and heat are at all equal to what is
produced here. This jelly-fish must be a deep-water species, as they have only been found during
a single autumn, and then only for a few days, when they seemed quite abundant."

f '

.-... :^,

Fin. lo<).—Ptychogtna " f>!nnulata"= P. laflea, after Haeckel, in "Report. Challenger Expedition, Deep-Sea Medusa.

v, velum; w, subumbrella; al, lips; urn, umbrella-margin; £r, stomach cavity; ob, marginal clubs; gs, ciliated
perradial cruciform furrow in the aboral wall of the central stomach; en, outer part of one of the radial-canals;
ck, distal funnel-like part of one of the radial-canals; j, gonad; u, umbrella; 1, tentacles.

— PTYCHOGENA.

217

Linko also considers Ptychogena to be a deep-sea form; as Browne, 1907, and Bigelow,
1909, state, however, we have no trustworthy evidence upon this point and it is not improb-
able that the medusa is merely a rare and somewhat variable Arctic species which lives
constantly at or near the surface.

Ptychogena longigona Maas.

Plychogcna longigona, MAAS, 1893, Ergeb. der Plankton Eiped., Bd. 2, K. c., pp. 64, 97, taf. 6, fign. 7-9.
Plychogena ( ?) longigona, BROWNE, 1907, Annals and Mag. Nat. Hist., scr. 7, vol. 10, p. 474.

Bell 25 mm. wide, flat, with firm gelatinous substance. About 100 tentacles very closely
set side by side so that they form an outer row around the margin, on the outer side of which
is a row of marginal clubs. Circular canal simple, but the 4 wide radial-canals have numerous,
indistinct, unbranched diverticula along their sides, the spaces between the lateral branches of
the canals being nearly rilled in by the gonads. Stomach shallow and 4-cornered and about
one-fourth as wide as the diameter of the bell. The mouth is a wide cruciform opening with
lips turned back as in Ptychogena, so that the water may enter the wide openings of the 4
radial-canals at the corners of the stomach. The gonads cover the entire lengths of the 4
radial-canals and their lateral surfaces give rise to hollow, evaginated, sac-like elevations or
papillae, which are neither so complex nor so regularly arranged as in Ptychogena. These

papillae occur along either side of each radial-
canal, but not in the median line. The mature
genital products are in the ectoderm. The
grooves of the 4 radial-canals extend along the
upper wall of the stomach and meet in the middle.
Color ( ?)

The species was found by the Plankton
expedition in the North Atlantic at a depth of 100
to 200 fathoms, but Maas does not give the exact
locality. As he states, it may prove to be only a
local variety of Ptychogena pinniilnin (Inctea).
It differs from Ptychogena I/itten, however, in
the simple side branches upon the radial-canals
and in that the gonads extend along the entire
length of the sides of the radial-canals, not being
restricted to their middle parts as in Ptychogfnu
lactea. The hollow elevations upon the gonads
are also simple and not so regularly arranged
as in P. lactea.

Medusae somewhat similar to P. longigona were found by the Plankton expedition near
the Cape Verde Islands.

Ptychogena? antarctica Browne.
Ptychogena antarctica, BROWNE, 1907, Annals and Mag. Nat. Hist., vol 20, ser. 7, p. 474.

The basal bulbs of the tentacles are laterally compressed and without ocelli. They resem-
ble those of P. lactea. Number of tentacles( ?) Sensory-clubs( ?) The 4 radial-canals have
wavy outlines, but no lateral branches. The wavy outlines of the radial-canals correspond with
those of the gonads, which are broad, lateral folds extending only part way along the radial-
canals. The stomach was lost in the single specimen studied by Browne. The tentacles are
red. Size of medusa( ?)

Found off Cape Andre, Victoria Land, Antarctic Ocean.

Ptychogena erythrogonon Bigelow.

Ptychogena erythrogonon, Bir.Et.ow, H. B., 1909, Bull. Mus. Comp.Zool. at Harvard College, vol. 57, p. 150, plates 5, 38, and 39.

This medusa is beautifully figured and well described by Bigelow.

Bell 38 mm. wide, 25 mm. high with very thick gelatinous substance and shallow bell-
cavity. 40 to 60 tentacles about one-fourth as long as bell-diameter and with tapering,
conical basal bulbs. There are only a very few cirri between the tentacles and these ter-

Fic. 110. — Ptychogena longigona, after Maas, in
Ergeb. Plankton Expedition.

218 MEDUSAE OF THE WORLD.

ininate in a knob-like cluster of nematocysts. Some of the medusae lack cirri and Bigelow
never saw more than 3 upon a single medusa. The cordyli or clubs upon the bell-
margin are more numerous than the tentacles. There are no lithocysts and no ocelli.
When young the outlines of the 4 broad radial-canals are plain. Later they develop short,
simple, lateral diverticula and then the gonads develop in the interspaces of these branches
close to the subumbrella as in P. longigona. The manubrium is quadrate basally and its
side walls are short and barrel-shaped with a widely open mouth.

Basal parts of radial-canals, manubrium, and tentacle bulbs are brilliant opaque brick-
red; gonads and outer parts of canal system and tentacles lighter. A pale colored line
extends along the central line of each radial-canal forming a cross at the middle of the
stomach. The entoderm cells contain red, yellow, and black granules, and the ectoderm
cells minute, red granules.

This medusa is one of the intermediate fauna found at depths of about 300 fathoms
in the Humboldt Current off the coast of Peru. Torrey describes another species of
Ptychogena from the California coast. (See appendix.)

Genus POLYORCHIS A. Agassiz, 1862.

Polyorchis, A. AGASSIZ in L. Agassiz's, 1861, Cent. Nat. Hist. U. S., vol. 4, p. 349; 1865, North Amer. Acal., p. 1 19. — HAECKEL,
1879, Syst. der Medus.n, p. 149. — MURBACH AND SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 174.

The type species is Polyorchis penicillata A. Agassiz, from the Pacific coast of the United
States.

GENERIC CHARACTERS.

Thaumantiadae with 4 radial-canals which give rise throughout their lengths to many
blindly ending side branches. Marginal ring-canal simple and bell-margin not divided into
lobes. Numerous sac-like, sausage-shaped gonads attached to the radial-canals and their side
branches. Stomach long, tubular, and with 4 simple lips.

Polyorchis penicillata A. Agassiz.

( ?) Medusa campanulata, DE CHAMISSO ET EVSENHARDT, 1820, Nova Acta Phys. Med. Leop. Car., tome I o, p. 359, plate 30, fig. I.

Meliccrtum pcniciltatum, ESCHSCHOLTZ, 1829, Syst. der Acalephen, p. 106, taf. 8, fig. 4.

Aglaura penicillata, DE BLAINVILLE, 1834, Man. d'Actinologie, p. 283, plate 33, fig. 4.

Polyorchis peniciUata, A. AGASSIZ in L. Agassiz's, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 349. — AGASSIZ, A., 1865, North Amer.
Acal., p. 119, figs. 179-183. — FEWKES, 1889, Amer. Naturalist, vol. 23, p. 593, figs. 3,4; 1889, Bull. Essex Inst. Salem,
vol. 21, No. 7, p. 103, plate 4, figs. 6, 7. — BANCROFT, 1904, Journ. Exper. Zool., Baltimore, vol. i, p. 289, 4 figs, (reac-
tions to galvanic currents). — BANCROFT, 1906, University of California Publications, Physio]., vol. 2, p. 43, 4 figs.

Polyorchis pinnatus+P. penicillatus, HAECKEL, 1879, Syst. der Medusen, pp. 149, 150, taf. 8, fig. 13.

Polyorchis — — , LOEB, 1906, Journ. Biological Chemistry, New York, vol. I, No. 6, p. 427 (physiological),

Polyorchh campanulata -+- P. penicillata, BEDOT, 1905, Revue Suisse de Zool., vol. 13, pp. 143, 144 (citation of all papers to l8<;o).

Polyorchis — — , LOEB, 1906, Dynamics of Living Matter, Columbia Univ. Biol. Series, No. 8, pp. 87, 91, 141.

Polyorchis — — , MACCALLUM, 1907, The Journal of Biol. Chemistry, vol. 2, No. 4, p. 385, New York.

Bell about 50 to 60 mm. high, 30 to 40 mm. wide, with a slight, solid, apical projection and
almost vertical, slightly bulging sides. Gelatinous substance quite thin. 36 to 40 tentacles
arise in a single row from the bell-margin. They are equal each to each and are longer than the
bell-height. Their basal bulbs are large, elongate, and spindle-shaped, and have dark-purple
pigment. There are no other marginal appendages. Velum narrow. Ring-canal simple
and unbranched, but the 4 radial-canals give rise each to 15 to 25 pairs of simple, short,
unbranched, blindly ending, lateral diverticula. Stomach prismatic with a very short peduncle,
and nearly as long as depth of bell-cavity. There are 4 short lips with sinuously folded mar-
gins. A row of 4 to 8 elongate, sac-like, sausage-shaped gonads arises from the sides of each of
the 4 radial-canals between the sides of the stomach and the feathered (branched) parts of the
canals, there being in all 16 to 32 of these suspended gonads.

Stomach, gonads, tentacle-bulbs, and radial-canals reddish-brown to purple.

Found on the Pacific coast of California to Washington, and as far westward as the
Hawaiian Islands.

Fewkes found the medusa abundant in bays along the California coast in March and
April. He states that when the medusa has 16 tentacles the 4 radial-canals were merely
beginning to exhibit notches along their sides and the gonads were beginning to appear at
the 4 corners of the stomach. When mature, the medusa has about 36 tentacles.

PLATE 26.

Figs, i and 2. LaoJicea? neptuna. Tortugas, Florida, July, 1898.
Fig. 3. Laodlceal neptuna. Part of the bell-margin.

Fig. 4. Staurophora rnertensii, half-grown medusa. Nahant, Massachu-
setts.
Fig. 5. Staurophora mertensii. Part of the bell-margin of the medusa shown

in figure 4.
Fig. 6. Staurophora mertensii, mature medusa, natural size. Agassiz

Laboratory, Newport, Rhode Island, June 9, 1895.
Fig. 7. Staurophora mertensii. Portion of the bell-margin of the medusa

shown in figure 6.
Fig. 8. Staurophora mertensii. Aboral view of the mid-region of one of the

radial-canals of the medusa shown in figure 6.
Fig. 9. Staurophora mertensii. Oral view of the mid-region of one of the

radial-canals of the medusa drawn in figure 6. This shows the

curtain-like folds of the gonads.
Fig. 10. StauroJiscus tetrastaurus, young medusa. Tortugas, Florida, July,

1898.
Fig. ii. StauroJiscus tetrastaurus. Oral view of female. Tortugas, Florida,

July, 1898.

See page 206 for description of figures I to 3.
^^^1 Drawn from life, by the author.

PLATE 26

. cr

/

11

'f

•/

"\ . . /•

v- . . .^

,fj '

\ • "r ' ' )

LEPTOMEDUS.E — POLYORCHIS, SPIROCODON.

219

Bancroft, 1904-06, carried out an interesting series of observations upon the galvanotropic
reactions of the medusa, showing; that the tentacles and manubrium respond definitely to the
current, even detached parts ot the tentacles reacting. The maximum response is at the anode
side in sea-water, whether acid or alkaline. The tentacles and manubrium turn and point
toward the cathode. The tentacles may be caused to contract slowly and irregularly in a con-
stant current. They contract when the current is made and relax when it is broken.

Loeb, 1906, finds that the magnesium of the sea-water serves to relax the muscles of the
bell, stomach, and tentacles of Pol\r>rcliis; and that the remaining salts of sea-water produce
a state of contraction, so that it we place the medusa in any solution resembling sea-water,
but lacking magnesium, pulsation is impossible and the tentacles and stomach are contracted.
If we add magnesium, however, pulsation is resumed, but this effect of magnesium can be
inhibited by addition of an equivalent amount of calcium or potassium.

The isolated center of the bell will not beat in sea-water or in a sugar solution, but
beats commence in these solutions if we add calcium, strontium, or barium chlorides. No

beats occur, however, if we add mag-
nesium chloride. The isolated center
will beat for 3 to 4 hours in any decalci-
fying salts, such as oxalates, fluorides,
oleates, citrates, etc. Acids cause the
center to beat, while alkalies inhibit
such beats.

The center does not beat until some
hours after being placed in NaCl; but
then rhythmical pulsations set in and
may last 2 or 3 days. But pulsation
may be at once produced in NaCl by
adding salts which precipitate calcium,
such as sodium citrate, tartrate, oroxa-
late. A trace of HC1 or CO2 added to
NaCl will also cause beats, as will pure
solutions of CaCl2, BaG2, or SnCl2.
Beats are inhibited more by !Mg than
by Ca. We have thus the \ anomaly
that contractions occur in an excess or
a deficiency of Ca.

Maccallum, 1907, finds that solutions of cascara sagrada, rhubarb, aloin,podophyllin,and
colocynth produce temporary pulsations in the isolated center of Polyorehis.

Polyorchis minuta Murbach and Shearer.

Polyorchis minula, MURBACH AND SHEARER, 1903, Proc. Zool. Soc. London, vol. z, p. 174, plate 19, fig. 3; plate 22, fig. i; 1902,
Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 72.

Bell 15 mm. high, 12 mm. broad. Truncated, oval, with thick walls; only 9 mm. wide at
velar margin. There are 55 or more tentacles. Yellow ocelli on outer sides of tentacle bases
on elevations. 4 narrow radial-canals with short, lateral diverticula. Stomach mounted on a
long peduncle extending to level of velar opening. 4 crenated lips. Gonads long, finger-shaped,
arising from the 4 radial-canals near base of peduncle.

Gonads and tentacles pale yellow in preserved specimens, other parts colorless.

Found in Puget Sound, British Columbia.

Genus SPIROCODON Haeckel, 1880.

SfirocoJon, HAF.CKEL, 1880, Syst. dcr Mcdusen, p. 636.

Goneomeandrus, KIRKPATRKK, 1903, Annals and Mag. Nat. Hist., scr. 7, vol. 12, p. 616.

The type species and only known form is Spirocodon saltatn\ ot the Inland Sea, Japan.
This was first described by Tilesius, 1818, as M,-Jus<i sallotrix. Haeckel, 1880, established
it as a type of the genus Spirocadon.

Fit:, ill. — Polyorchis ptnicillala, after Fewkes, in
American Naturalist.

220

MEDUSAE OF THE WORLD.

GENERIC CHARACTERS.

Thaumantiadae with 4 radial-canals and a ring-canal, all of which give off blindly ending,
dendritic side branches. Bell-margin cleft into lappets. Stomach mounted upon a peduncle,
and with 4 lips. Gonads sac-like, complexly folded, and developed upon the radial-canals
adjacent to sides of stomach.

This genus resembles Polvorchis, but ring-canal gives off blindly ending, centripetal
branches, and margin is cleft into lappets.

Spirocodon saltatrix Haeckel.

Medusa saltatrix, TILESIUS, 1818, Mem. Acad. St. Petersbourg, tome 6, p. 554, taf. 18.
Polyorcliis saltairix= Spirocodon saltatrix, HAECKEL, 1880, Syst. der Medusen, p. 636.

Spirocodon sa/tatrix, GOETTE, 1886, Sitzungsber. kgl. Preussisch. Akad. Wissenschaft. zu Berlin, Jahrg., 1886, p. 832.
Goneomeandrus chryiostephanus, KIRKPATRICK, 1903, Annals Mag. Nat. Hist., ser 7, vol. 12, p. 616, plate 33, figs. 1-4 (this is
Spirocodon saltatrix Haeckel; see Kirkpatrick, 1904, Annals and Mag. Nat. Hist., ser. 7, vol. 13, p. 80).

Bell with octagonal sides and dome-like, rounded top, angles of side-walls radial and in-
terradial in position, 30 to 60 mm. wide, 50 to 80 mm. high. 8 adradial, semicircular, marginal
lappets. The clefts between the lappets pass obliquely upward and outward from below and
extend much higher on the exumbrella than on the subumbrella side, so that only the outer
third of each lappet is free. The lappets alternate in position with the angles of the sides of
the bell. Velum quite broad, its inner edge circular.

112.

"3-

FIG. 112. — Spirocodon sallatrix, after Kirkpatrick, in Annals and Mag. Nat. Hist.

FIG. 113. — Cannota dodccantha, after Agassiz and Mayer, in Bull. Museum Comp. Zool. at Harvard College.

There are about 40 to 80 tentacles on each of the 8 lappets, 326 to 640 in all. These ten-
tacles arise in a single row from the margin, and are about one-third as long as bell-diameter.
Their bases are hollow and narrower at margin than at some distance out. There is an
orange-brown ocellus above the base of each tentacle and there are no other marginal sense-
organs. There are 4 radial-canals and a ring-canal which extends around the margins of the
lappets. All of these canals give off numerous blindly ending, dendritic side branches, which
do not anastomose. 4 main centripetal vessels arise from the interradial angles of the mar-
ginal ring-canal and smaller ones arise from the ring-canal in adradial and other positions.

The stomach is set upon a short, gelatinous peduncle which is one-thud as wide as the bell
itself and as long as wide; it is cruciform in cross-section, 4-sided, and about half as long as

LKPTOMKDfS.K— SI'IKorniNiX, CANNOTA, ' 1 VIKHIA. 221

bell-height. There are 4 slightly recurved lips with sinuously folded margins. The gonads
are complexly folded, swollen, projecting masses on proximal parts of the 4 radial-canals, upon
the peduncle adjacent to the stomach. The bell is colorless, gonads pale yellow,
orange to brown. Found at Nagasaki, Inland Sea, Japan.

Genus CANNOTA Haeckel, 1879.

n, p. 151.

sber. math.-phys. Klasse kgl. Bayer. Aka

The type species is Cannota doJciiintlia Haeckel, from New Guinea, tropical Pacific.

Canrtota, HAECKEL, 1879, Syst. dcr Medusen, p. 151.

Staurodhcu! (in part), MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer. Akad. Wisscnschafi., Bd. 34, p. 440

GENERIC CHARACTERS.

Thaumantiadie with 4 main radial-canals, each giving rise to 2 simple, unbranched
side branches which, together with the main radial-canals, connect with ring-canal at margin.
Gonads upon the 4 main canals and side branches. Marginal clubs and cirri may or may not
be present. 12 or more tentacles.

This genus is closely related to St/uuoJmus, but is distinguished by the fact that the side
branches of the radial-canals join the marginal ring-canal, whereas in StauroJiscus they end
blindly.

Maas, 1904, would regard StiiiirnJi.^-ns as a young stage of Haeckel 's ('.unnntn. but nix-
studies of numerous specimens of Xtiiii>-rnlis?ns at Tortugas, Florida, convince me that the
medusae become mature with blindly ending side branches upon the 4 main radial-canals.

Cannota dodecantha Haeckel.

Cannota dodecantha, HAECKEL, 1879 Syst. der MeJuscn, p. 151.

Staurodhcus nigricans, AGASSIZ, A., AND MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 164, plate 4,
figs. II, ll. — BROWNE, 1907, Annals and Mag. Nat. Hist., vol. 20, ser. 7, p. 475.

Bell thin and flexible, 14 mm. in diameter. 12 short tentacles with well-developed basal
bulbs. The 4 tentacles at the bases of the 4 main canals are about twice as long as the 8 which
arise from the ends of the lateral branches. The longest tentacles are, however, only about one-
sixth as long as bell-diameter. Between each successive pair of tentacles there are J small
protuberances upon the bell-margin, each one of which bears a sensory-club. Velum narrow.
Each of the 4 radial-canals gives rise to a pair of opposed lateral branches and thus 12
canals reach the circular vessel 30° apart. The gonads are developed upon these 12 trident-
like terminal canals. Manubnum is \et\ short, mouth-opening wide. Gelatinous substance
and entodermal lamella of bell of a brownish tinge. Genital organs, basal bulbs of tentacles,
and manubrium are light-sepia in color. Bulbs of sensory-clubs are dark-brown.

This medusa was found near Suva Harbor, Fiji Islands, South Pacific, in December, 1897.
Haeckel describes a small specimen 4 mm. wide from New Guinea. (See fig. 1 13.)

Cur

Cuv

Cui
Cm
Cm

Genus CUVIERIA Pe"ron, 1807.

eria (in botany), KOBREUTER, 1802; DE CANDOLLE, 1807.

eria (medusa), PERON, 1807, Voyage aux terres australcs, plate 30, figs. 2, 2a.— AC.ASSIZ, L., 1862, Com. Nat. Hist. V. S.,

vol. 4, p. 345.

eria (echinoderm), PERON, 1817.
eria (crustacean), DESMAREST, 1825.
eria (mollusk). RANG, 1828.

Berenix, PERON ET LESUEUR, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 327.

Berenice, OKEN, 1815, Lehrbuch der Naturgcsch., Teil 3, Zoul., p. 1 14.— ESCHSCHOLTZ, 1829, Svsl. der Acalrphrn, p. 120. —
MILNE-EDWARDS, 1849, Cuvier's Regne animal illvistrr. /.ii»|>h.. plate ;',, fig. I, la.— HMC KM . iS-9. Sv-t. .|rr Mrdusen,
p. 152. — MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer. U-.i.l. \\ i ^ MM h.ifi., B.I. -,4, p. 440. — BF.DOT, 1901,
Revue Suisse de Zool., tome 9, p. 482; Ibid., 1905, tome i-,, p. i •, i (list of all references from 1807-1850).

As is apparent from the above, the name C.ir, i,-im was first used in botany in 1802, and in
zoology for a medusa in 1807. Its previous use in botany does not inuiK-u- \\ith its use in
zoology. The type species is Cuvicria carisockroma of Peron from the tropical Atlantic.
18

222

MEDUS.E OF THE WORLD.

GENERIC CHARACTERS.

Thaumantiadae with 4 principal radial-canals which branch, and the branches again
branch, and all unite with the circular vessel. The gonads are developed upon the outermost
branches of the radial-canals near the ring-canal. Tentacles numerous. There may or may
not be marginal clubs or cirri. Development unknown.

Modern authors have commonly followed Peron and Lesueur, 1809, and Eschscholtz,
1829, in calling this genus Berenice, but if we follow the rule of precedence the name Cuviena
must stand. It is rarelv a pleasant duty to revive an unfamiliar name to supplant one well

known to literature, but happily in this case the
name of one whose memory we all delight to honor
may once more be associated with a genus in that
branch of science which his labors ably advanced.

Cuvieria carisochroma P£ron.

Cuvieria carisochroma, PERON, 1807, Voyage aux terres australes,
plate 30, figs. 2, 2a.

."Equorea euchroma and fequorea rosea, DE LAMARCK, 1817, Anim.
sans vertebres, tome 2, p. 497.

Berenice rosea, ESCHSCHOLTZ, 1819, Syst. der Acalephen, p. 120. —
HAECKEL, 1879, Syst. der Medusen, p. 153. — BEDOT, 1901,
Revue Suisse de Zool., tome 9, p. 482; Ibid., 1905, tome 13,
p. 131 (list of papers to 1850).

(?) Berenice capillata, HAECKEL, 1879. Syst. der Medusen, p. 154,
taf. 9, fig. 5.

According to Peron, bell is flat and shield-
shaped, about 50 mm. wide and 10 mm. high.
There are 50 to 60 very long thin tentacles with
very large, hemispherical basal bulbs. Each of the
4 principal radial-canals gives rise to two opposed
side branches, and these side branches as well as
the principal canals give rise to 3 to 5 terminal
branches which reach the circular canal. Thus 50
to 60 canals reach the circular vessel. The 50 to 60
gonads are developed on terminal branches of
canals. Bell, gonads, and tentacles rose-red, and
tentacle-bulbs yellow. This species is found in the
tropical Atlantic.

I am inclined to regard Haeckel's Berenice
capillatij as being identical with this medusa.
According to Haeckel the bell in B. " capillala" is
hemispherical, 20 mm. wide, 100 to 120 long ten-
tacles, with well-developed globular, spindle-shaped
basal bulbs. There are no marginal clubs or cirri.

The 4 main radial-canals and their 2 side
branches each and all give rise to 4 to 8 terminal
branches, so that 50 to 100 vessels reach the
marginal ring-canal. Small, spindle-shaped gonads
are developed upon the terminal branches of the radial-canals near the ring-canal. Stomach
small, with 4 simple, recurved lips. Color (?)
Cape Verde Islands, Atlantic Ocean.

Cuvieria huxleyi.

Berenice huxleyi, HAECKEL, 1879, Syst. der Medusen, p. 154, taf. 9, fig. 4.

Bell flat, 16 mm. wide, 0.4 mm. high. 16 long tentacles with globular to spindle-shaped
basal bulbs with ocelli. 80 to 100 marginal clubs, 2 to 6 between each successive pair of ten-
tacles. Stomach small, flat, with 4 simple, short, cruciform lips. The 4 main radial-canals

FIG. 114. — Cuvieria "capillnla"= C. carisochroma,
after Haeckel, 1879.

LKITO.MKDrs.K -< T VIKHIA. DK 'H< )T< iM I \.

223

each give rise at their middle points to 2 opposite side branches. These main canals and their
2 branches each end in 5 to 7 terminal branches, and thus about 70 vessels reach the ring-
canal. Gonads spindle-shaped, on terminal branches of the canals near the ring-canal.
Color (?) Found off Azores, Atlantic Ocean. Distinguished by its large number of
marginal clubs and few tentacles. Is it only a variety of ('111-1,11/1 ciirisochrntnii 1'eron ( ?)•

Genus DICHOTOMIA Brooks, 1903.

Dichotomia, BROOKS, 1903, Proc. American Philosophical Society, Philadelphia, vol. 42, p. i i . - MA*S, 1904, Siuungsber. math.-
phys. Klasse kgl. Bayer. Akad. dcr Wisscnschaften, Bd. 34, Heft 3.

CKNKRR CHARACTERS.

Berenicidae with 4 main radial-canals which divide dichotomously 2 or more times and
open into the circular vessel by 16, 32, or more distal branches. Tentacles of t\\<> suits
hollow ones and solid ones — all arising from bell-margin. Mouth simple without prominent
lips. The gonad encircles the stomach and extends outward over the radial-canals and their
branches.

115.

116.

Fie,. 115. — Curifria huxltyi, after Haeckel, 1879.

Flc;. lib.— Dichotomia cannoides, after Brooks, in Proc. American Philosoph. Soc., vol. 42.

A, young medusa. B, adult medusa. C, model showing arrangement of radial-canals.

The type species is Dicliutniniu < annoides, described by Brooks from the Bahama Islands.
This is the only known species.

Dichotomia cannoides Brooks.

Dichotomia cannoides, BROOKS, 1903, Proc. American Philosophical Soc. Philadelphia, vol. 42, No. 172, p. 12, plate i, 3 figs. —
MAAS, 1904, Sitzungsbcr. math.-phys. Klasse kgl. Bayer. Akad. der \VisM-nschaft., Bd. 34, p. 438.

Bell about 8 mm. high and 6 mm. wide, subcylindrical, and with conical apex. The 4
main radial-canals do not arise independently and directly from aboral end of stomach, but in
pairs from the ends of a short transverse canal, so that the only planes which divide the medusa
into symmetrical halves are the two primary interradial planes, \\hen thus divided each halt
is bilaterally symmetrical, and the halves are reversed copies of each other (fig. i i<' i . The
4 primary radial-canals branch dichotomously at least 3 times, so that 32 terminal branches

224 MKIH'S.K OF THK WORLD.

reach the circular vessel. The radial-canals of older medusae may branch more profusely.
The subumhrella consists of two strongly contrasted regions: An upper opaque part, nearly
hemispherical, contains the arches formed by the gonad, which spans space between proximal
pairs of radial-canals and stomach in a system of groined arches. The lower half of the
subumhrella is transparent and without these arches.

There are 16 long, hollow, contractile tentacles with tightly coiled tips. In the young
medusa, with only i" terminal radial-canals, these tentacles arise from margin at base of each
canal; but in older medusae, with 32 terminal branches of radial-canals, the hollow tentacles
are in the radii of dichotomy. In addition to the long, hollow tentacles, there are a number of
short, stiff, solid tentacles, which remind one of the solid tentacles of Geryonidae, and are
carried turned upward from bell-margin. These short, stiff tentacles are equal in number
to the terminal branches of radial-canals.

There are no marginal sense-organs, clubs, or cirri. The manubnum is a truncated cone,
extending about two-thirds the distance from inner apex of bell-cavity to margin.

The gonads and manubrium in old medusae are opaque white. The radial-canals, cir-
cular vessel, and axes of hollow tentacles in young medusae are colored by brownish to orange
pigment-granules. The bell, subumbrella, and tentacles are nearly colorless in old medusas.

This interesting medusa was found by Professor Brooks at Nassau, Bimini, and Green
Turtle Key among the Bahama Islands, and was described by him in detail.

Genus DIPLEUROSOMA Axel Boeck, 1866, sens, emend.

Difleurosoma, AxFLBoECK, 1866, Forenings Vidensk. Meddelclser, Nos. 10, 1 1, pp. 131,136. — HAECKEL, i879,Syst. der Medusen,
pp. 155, 636. — AOASSIZ A. AND MAYER, 1902, Mem. Mas. Comp. Zool. at Harvard College, vol. 26, p. 148. — BROWNE,
1898, Proc. Zool. Soc. London for 1897, p. 826; 1900, Proc. Royal Irish Acad., ser. 3, vol. 5, p. 715.

Telracannota, MATER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 46; 1904, Mem. Nat. Sci. Brooklyn Inst .
Mus., vol. i, p. 12.

Tetracannota + Dipleurosoma, MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer. Akad. der Wissenschaften, Bd. 34, pp.
440, 441.

Ametrangia, ALLMAN, 1873, Nature, vol. 9, p. 73.

The type species is Dipleurosoma t \picnm Axel Boeck, 1866, from the coasts of Norway,
Newfoundland, and northern Atlantic generally.

GENERIC CHARACTERS.

Thaumantiadae with 3 or more main radial-canals, some or all of which branch irregularly
(not regularly and dichotomously) and connect with the circular vessel. Gonads are upon the
radial-canals adjacent to manubrium. Tentacles numerous, and marginal clubs, cirri, and
ocelli may be present. The hydroid appears to be CuspiJelln.

The radial-canals vary greatly both in number and arrangement in /). t\t>icntn, and are
best described by Browne, 1900. This extreme variability has caused the species to he
described under various names, fudging from the irregular arrangement in the single specimen
of D. pacifica found by Agassiz and Mayer, near Tahiti, the canals may be equally variable.
In D. collapsa they appear to be more regular, mature specimens having 16 canals arranged in
4 groups of 4 canals each.

Dipleurosoma typicum Axel Boeck.

Dipleurosoma typitum, AXEL BOECK, 1866, Forenings Vidensk. Meddelelser, Nos. lo-II, p. 131, fign. 1-3; Ibid., D. stuvitzi, pp.

136, 137, figs. 1-4. — BROWNE, 1900, Proc. Royal Irish Acad., ser. 3, vol. 5, pp. 696, 715, plates 20, 21. — MAAS, 1904,

Sitzungsber. math.-phys. Klasse kgl. Baver. Akad. der Wissenschaft., Bd. 34, p. 441.
Ameirangia htmhpharica, ALLMAN, 1873, Nature, vol. 9, p. 73.

Difleurosoma tyficum+amfliithectum+ irregulare, HAECKEL, 1879, Syst. der Medusen, pp. 155, 636, taf. 9, fig. 9.
Dipleurosoma hemisph:rrica,a\DDON, 1885, Proc. Roy. Irish Acad., ser. 2, vol. 4, p. 526. — BROWNE, 1898, Proc. Zool. Soc. London,

for 1897, p. 826, figs. 10-12.

Bell usually flatter than a hemisphere and about 15 mm. wide. Marginal tentacles very
numerous, more than 100. Each tentacle has a bulbous base with an ocellus upon its inner
side. There appear to be no marginal clubs in D. tvpicum, but these occur in D. pacifica of the
tropical Pacific. Velum well developed. Stomach is flat and watch-glass-shaped, with an
irregular outline, and there are 4 lips. In 217 specimens studied bv Browne from Valencia

I.I I'l' 'MUM > I! Diri.KI IdisnM v.

Harbor, Ireland, the number of radial-canals arising fiom the periphery of tin stomach
ranged troni 5 to |H, 43 of the medusae hav ing .y canals. Canals exceedingU irregular in position
and in mode of branching, so that it is impossible to define a normal t\pe. but rhe\ appear nor-
mally to connect with the circular \essel. The Bounds are developed upon radial-canals adja-
cent to the stomach and may be upon all or on onl) a few of the canals. Tin \ van in number
from i to 12, and 5 is the most frequent number. The females are in excess of the males in
the proportion of about 4 to 3. The ova remain attached to the ovary until the planula sta»e
is reached. They were partially reared by the Misses Delap, and developed into hydioids
apparently belonging to the genus Cuspidella isee Browne. 1900, p. 696).

The gonads, stomach, and tentacle-bulbs are said to be brownish (Haeckel. iS;

The changes which the medusa undergoes in its growth are unkno\\n, but Browne be-
lieves that the stomach is primarily regular in shape and that it becomes irregularly elongated
into lobes by development of the radial-canals.

118.

Fu;. 117. — Dipleurosoma ("amphitheetum"'), after H.irckcl, 1879.
l'i'.. iiS. -Dipleurosoma pticificti, atlt-r N I . in

\lu -rum Cornp. Znol. at Harvard C<>llri;<..

D. tvpii-iuii of the Ninth Atlantic may prove to be identical with /). [>iii'i/ii-a from Tahiti,
South Pacific. The only distinctions between the North Atlantic and the tropical Pacific
species are their wide geographical separation and the presence of' maiginal clubs in I).
piififiin. Dtpleurosoma tvp/fiiiii has been taken ott the northern coasts of Europe and <>rl
Newfoundland.

Dipleurosoma pacifica Agassiz and Mayer.

Diplcurosoma pacifica, Ac, AS si z A., AND MAYI K, 1902. Me in. Muv. Com p. /.""I . at H.n uir^l (.'"II ., \"\ . ;l>. p. i 48, plate j, figs. 13, 14.

Bell very flat, about j times as broad as high, 2} mm. in diameter. Surface of exumbrella
reticulated by a regular system of hexagonal elevations bounded b\ shallow turn>\\s. About
100 short tentacles with narrow, bulbular bases. Kntodermal core of these tentacles is solid
and composed of chordate cells. At the bases of most tentacles upon the lower (subumbrella)
side, there is a single black ocellus. Slender, club-shaped, sensory bodies aie scattered at fre-
quent and irregular intervals between the tentacles. Entodermal cores of these bodies air solid
and very similar to those of the tentacles. No concretions. Velum narrow. <> radial-canals
arise in groups of 3 from diametrically opposite sides of the stomach. In the single specimen
examined, 2 of these canals bifurcate, and 8 canals thus reach the circular vessel. The <;onads
are linear and occupy the middle regions of the S canals. Their surfaces are slighrlv papillate
and the ova stand out as hemispherical protuberances. Stomach elongated in the direction of
the two opposed groups of radial-canals. Manubrium very short, 6 small, slightly folded lips.

Tentacle-bulbs, genital organs, and manubrium milky white, all other parts of the medusa
being transparent. A single specimen was obtained in a surface haul about 50 miles ninth of
Tahiti. Smith Pacific.

220 MEDUSA OF THE WORLD.

Dipleurosoma is exceedingly variable in the number and arrangement of its radial-canals.
The Pacific species is distinguished by having marginal clubs between its tentacles, these being
absent ( ?) in the Atlantic Dipleurosoma typicnm.

Dipleurosoma collapsa Mayer.

Plate 27, figs. I to 3, and 7.

Tftracannolatolla/>sa,MA\T.tt, 1900, Bull. Mus.Comp.Zool. at Harvard College, vol. 37, p. 46, plates 7,8, figs. 14-16; Memoirs Nat.
Sci. Mus. Brooklyn Inst. Arts and Sci., vol. i, No. i, p. 12, plate 4, fig. 32. — MAAS, 1904, Sitzungsber. math.-phys. Klasse
lier kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 440.

Adult nii-Jusa (plate 27, fig. 3). — Bell J mm. in diameter and about as high as broad.
Tup dome-shaped, side walls vertical. 16 well-developed tentacles are carried tightly coiled
in close helices, and besides these there are 1 12 very small rudimentary tentacles. Dark-brown
entodermal pigment-spots are found within the base ot each tentacle. There are 16 radial-
canals, arranged in 4 groups of 4 each (plate 27, fig. 7). The gonads are found in proximal
parts of the 16 radial-canals very near the point where they branch off from the stomach.
Peduncle of manubrium wide and prominent. The mouth has 8 slightly crenated lips.
Entoderm of manubrium in some specimens is green, in others it is pearly-white or yellowish.
Entodermal pigment-spots at bases of tentacles are dark-brown.

Stugi-s in development. — The youngest medusa observed had a bell i mm. in diameter
(plate 27, fig. i ). It had 4 slender radial-canals and 24 tentacles, 4 well-developed and 20
rudimentary. Velum prominent; 4 lips, and as yet no peduncle. No trace of genital organs.
In the next older stage (plate 27, fig. 2), we find 4 new radial-canals beginning to develop. A
still older stage (plate 27, fig. 7) has 16 radial-canals and 8 lips, but as yet no peduncle and no
trace of gonads, nor have radial tubes grouped themselves into 4 bundles as in the adult.

This medusa is very common at the Tortugas and Bahamas in June, and ample oppor-
tunity for observing its transformation was afforded. It exhibits the curious habit of collap-
sing into an almost shapeless mass, in which condition it may remain tor several hours and
then "straighten out" and swim about in excellent condition.

Fewkes, 1883 ("On a Few Medusa? from the Bermudas," Bull. Mus. Comp. Zool.,
vol. ii, No. 3, figs. 7, 70), has evidently figured the young of this species under the name of
"Larva of an unknown Tubularian."

Dipleurosoma ochracea sp. nov.
Plate 29, figs. I and 2.

Immature medusa. — Bell thin-walled, somewhat flatter than a hemisphere and 8 mm. in
diameter. About 12 to 16 well-developed tentacles and about 35 immature or rudimentary
tentacle-bulbs. The well-developed tentacles are in the radii of radial-canals. They have
hollow, conical, basal bulbs and filiform, contractile, lash-like shafts, which are about as long
as the bell-radius. Each tentacle-bulb is flanked by a pair of large, sausage-shaped, marginal
clubs, and in addition to these there is a single sausage-shaped diverticula from the ring-canal
on the subumbrella side above each tentacle-bulb. There are no marginal cirri or append-
ages other than those described above. Velum very wide. Central stomach is flat and about
one-fifth as wide as bell-diameter. About 6 to q radial-canals arise in a very irregular man-
ner from margin of stomach; some of these give rise to side branches. Manubrium very
shallow, 6 or more very irregularly arranged, folded lips with plain margins. No gonads are
observed, though they may develop later upon the broad, expanded middle regions of the
radial-canals. Stomach, radial-canals, and marginal clubs of this medusa are crowded with
masses of yellow algae, or Philozoon, apparently similar to that which infests the medusae of
/ elella. These plant-cells develop between the cells of the medusa and give a dense, opaque,
brilliant yellow coloration to the marginal clubs, and in a lesser degree to the stomach and
canals. All other parts are colorless.

The medusae are remarkable for their very active, incessant pulsation.

Large swarms of this medusa appeared at Tortugas, Florida, between July 5 and 24, 1907,
but no mature specimens could be found.

PLATE 27.

Fig. I. Dipleurosoma collapsa, young medusa with 4 simple radial-canals.

Tortugas, Florida. June, 1897.
Fig. 2. Dipleurosoma collapsa, half-grown medusa with 16 simple, separate,

radial-canals.
Fig. 3. Dipleurosoma collapsa, mature female, with 16 radial-canals grouped

into clusters of 4 canals each. Tortugas, Florida, July, 1898.
Figs. 4 and 5. N etocertoides brachiatum. Successive stages in the growth of

the medusa. Tortugas, Florida; and Nassau, Bahamas, July,

1898 and 1903.

Fig. 6. N etocertoides braclnatum. Oral view of medusa shown in figure 4.
Fig. 7. Dipleurosoma collapsa. Aboral view of the stomach showing the

grouping of the radial-canals into 4 radial clusters ot 4 canals

each. Tortugas, Florida.

Fig. 8. Turris superba. Aboral view of the stomach showing the inter-
radial positions of the gonads.

See page 126 for description of figure 8.
Drawn from lite, by the author.

PLATE 27

— DIPLKUiOSMM \.

227

Dipleurosoma brooksii sp. nov.

This form is described from drawings made by the late Prof. William K. Brooks, and
found after his death among his unpublished figures. The drawings were generously pre-
sented to the author by the Department of Biology of the Johns Hopkins University for
reproduction in this work.

Bell flatter than a hemisphere, evenly rounded, thin walled. Manubrium small, short,
and 8-sided. 8 simple lips, 4 long and 4 short. 8 radial-canals arise from the stomach,
the 4 in the radii of the long lips are trident-shaped, each giving rise to 2 side branches.
The 4 canals in the radii of the short lips are simple. Thus if> terminal branches reach
the circular vessel. In the specimen figured by Professor Brooks only 14 canals joined with

Fir,. Il8a. — Dipleurosoma brooksii, sp. nov. A, oral view; n, siilc view
with tentacles cut off short; c, oral view of lips.
Drawn from life by Professor Brooks.

the circular vessel, for two of the side branches of the main radial-canals tailed to develop.
There appear to be typically 16 tentacles, one at the base of each terminal branch of the
radial-canals, although in two intervals there are small additional tentacles thus giving 18
tentacles in the specimen figured by Professor Brooks. The tentacles have long, conical,
tapering basal bulbs, which are hollow and very long, lash-like shafts, longer than the bell-
diameter and very flexible. No ocelli are figured by Professor Brooks. The gonads are
upon the sides of the 16 radial-canals, not touching the ring-canal but extending from the
sides of the stomach outward. The velum is a wide, annular diaphragm. The size and
color of the medusa can not be determined from Professor Brooks's drawings. The
medusa was found at Nassau, Bahama Islands.

228

OF THE WORLD.

Were it not for the absence of ocelli I would be inclined to regard this medusa as a
hypertrophic specimen of Staurodiscus tetrastaurus in which the side branches of the 4
primary radial-canals had reached the circular vessel and 4 intermediate canals had
developed. I have, however, never observed such a condition among many hundreds of
apparently mature medusas of S. tetrastaurus found at Tortugas, Florida ; and it seems
probable therefore that Professor Brooks's medusa is a new species which may have been
derived from a Staurodiscus-like ancestor.

Genus TOXORCHIS Haeckel, 1879, sens. ampl.

Toxorchii + Cladocanna, HAECKEL, 1879, Syst. der Medusen, pp. 156, 160.

Toxorcliis, MAAS, 1904, Sitzungsber. math.-phys. Klasse kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 441.

Haeckel, 1879, founded this genus for Toxorchis arcuatus of the Canary Islands, Atlantic
Ocean; but the first known species is T. thalassina=Berenix tluilnssina Peron and Lesueur,
1809, from Northern Australia.

GENERIC CHARACTERS.

Thaumantiadae with 4, 6, or more main radial-canals which branch regularly and dichot-
omously one or more times. The gonads are upon the outermost branches of the radial-canals,
near the ring-canal. Numerous tentacles, cirri, and marginal clubs. Hydroid unknown.

Haeckel restricts Toxorchis to include medusae having radial-canals branched once
dichotomously, while his Cladocanna applies to medusae having radial-canals with 2 or more
dichotomous branchings.

Toxorchis thalassina Maas.

Berenix lhalassina, PERON ET LESUEUR, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 327.

Berenice euchroma, DE BLAINVILLE, 1834, ManXiel d'Actinologie, p. 276, plate 32, fig. I.

Cm'ieria euchroma, LESUEUR, 1839, m Cuvier's Le Regne animal, plate 53, fig. 2.

Cladocanna thalassina+ C. pol\clada, HAECKEL, 1879, Syst. der Medusen, pp. 160, 161.

Cladocanna polyclaiia, YON LENDENFELD, 1884, Proc. Linnean Soc. N. S. Wales, vol. 9, p. 581.

Toxorchis thaiassiria, MAAS, 1904, Sitzungsber. math.-phys. Kla&se kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 441.

Cladocanna thalassina, BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 482; Ibid., 1905, tome 13, p. 132 (list of papers to 1850).

Bell flatter than a hemisphere to hemispherical, about 50 mm. wide. About 100 or more
long tentacles with marginal clubs and cirri between them. Stomach shallow, with 6 wide lips.
6 wide radial-canals arise from stomach. For one-third the distance between stomach and
margin these canals are simple and unbranched, but outwardly they branch dichotomously
and somewhat irregularly 3 to 4 times, so that there are about 100 terminal branches which
reach the circular canal at the margin. The sac-like, spindle-shaped, folded gonads are
developed upon the outermost branches of canals. The entire medusa is light sea-green.

Found off the coasts of New Guinea and
Northern Australia. Development unknown.

Toxorchis arcuatus Haeckel.

Toxorchis arcuatus, HAECKEL, 1879, Syst. der Medusen, p. 157,
taf. 9, figs. 6-8.

Bell flat to hemispherical, 6 mm. wide.
Stomach small and flat, with 6 much crinkled,
lanceolate lips. 6 main, wide radial-canals
which fork at their middle points, so that 12
canals, 30° apart, reach the circular vessel at
bell-margin. There are 24 long tentacles with
bulbous bases, each bearing a reddish-brown
ocellus. Numerous clubs and cirri on bell-
margin between the tentacles.

The gonads are developed upon the edges
of the forked ends of radial-canals, there being
iatu*, after Haeckel, 1879. a crescentic gonad upon the concave side of the

crotch of each fork, and a pair of linear gonads upon the convex sides of the forks. There
are thus 3 gonads upon each main canal at its forking, 18 gonads in all.

PLATE 28.
Fig. i. Toxorchis kellneri. Side view of female medusa. Tortugas, Florida,

July 8, 1907.

Fig. 2. Toxorchis kellneri. Oral view of the medusa.
Fig. 3. Turris superba (a variety of T. pileata). Tortugas, Florida; July

4, 1907. Mature male medusa.
Fig. 4. Turris superba (variety of T. pileata). Side view of manubrium

showing the foldings of the interradial gonads.

See page 126 for description of figures 3 and 4.
Drawn from life, by the author.

PLATE 29.

Fig. i. Dipleurosoma ochracea. Side view of an immature medusa.

Fig. 2. Dipleurosoma ochracea. Oral view of an immature medusa.

Fig. 3. Stomotoca pteropliylln. Enlarged view of the medusa, showing the
bell contracted so as to reduce the length of the peduncle. Nassau
Harbor, Bahamas, April 20, 1907.

Fig. 4. Stomotoca pterophylla. Side view of the manubrium of a mature
male. Off Miami, Florida, in the Gulf Stream.

Fig. 5. Stomotoca pterophylla. View of velum showing radial and circum-
ferential muscle-fibers.

Drawn from life, by the author.

See page 113 for description of figures 3 to 5; page 226 for descrip-
tions of figures i and 2.

PLATE 29

LEPTOMEDUS.E — TOXORCHIS, NETOCEKTi )1 in >. --".'

Stomach, lips, gonads, and tentacle-bulbs are yellowish-brown, speckled with dark-red.
Exumhrella besprinkled with reddish-brown spots.

Found bv Haeckel at the Canar}- Islands, Atlantic Ocean, in January. 1867.

Toxorchis kellneri sp. nov.
Plate 28, figs, i an>l j.

This medusa is named in honor of the author's assistant, Mr. Carl Kellner. whose copies
of figures ot medusae by previous authors appear as text-figures in tins work.

Bell about 15 mm. wide, with moderately thin walls, thickest at center ot" bell and thin
at edges. When in a normal resting state the bell is somewhat higher than a hemisphere,
but when expanded it is slightly Hatter than a hemisphere. There are about $2 tentacles
with long, conical, tapering, hollow basal bulbs. The filiform shafts ot these tentacles are
highly contractile and normally i to 4. times as long as the bell-diameter, but may be con-
tracted so as to become about as short as the bell-radius. There are about 50 small clubs
upon the bell-margin between the tentacles; but no coiled cirri nor lithocysts. 1 here is a black
entodermal ocellus within ring-canal at base ot" each tentacle and marginal club.

Velum well-developed, ring-canal narrow. Stomach about one-fourth as wide as bell-
diameter. 8 flat, wide radial-canals arise trom margin ol stomach, 45° apart. A short dis-
tance beyond the margin ot the stomach most ot these 8 primary radial-canals bifurcate.
Thus in the specimen here fimired n ot the 8 primary radial-canals are bifurcated and thus
14 slender radial-canals reached the marginal ring-canal.

The manubrium is shallow, only about one-fourth as long as the depth ot the hell-cavity,
and there are 8 lanceolate lips with folded, curtain-like margins. These lips are in the radii
of the 8 main radial-canals. The gonads are developed upon the sides of the narrow outer
branches of the radial-canals and do not touch the ring-canal.

Central stomach pink; gonads and tentacle-bulbs brownish-yellow; entodermal ocelli
very dark brown, almost black.

Several specimens of this medusa were found in a surface tow at Tortugas. Florida,
on July 8, igo/. The figures represent a female.

This species differs from Haeckel 's T. uriiintits ot the Canary Islands in its higher bell,
more numerous and more irregularly branching radial-canals, in the absence of marginal
cirri, and in color. Unfortunately, Haeckel had but one specimen, and I have seen only three,
and it is possible that future studies may show that the Canary Island medusa is identical
with the Tortugas form.

Genus NETOCERTOIDES Mayer, 1900.

NcloctrtoiJes, MAYER, 1900, Bull. Mus.Comp. Zool. at Harvard College, vol. 37, p. 45.— MAAS, 1904, Sitzungsber. math.-phys.
Klasse kgl. Bayer. Akad. Wissenschaft., Bd. 34, p. 437.

GENERIC CHARACTERS.

Berenicidae with 8 main radial-canals, which bifurcate so that 16 canals reach the circular
vessel at bell-margin. There are 16 long, hollow, tapering tentacles, one at the end of each
radial-canal, and 16 to 25 short, cirrus-like tentacles, one or t\\o between each pair ol long
tentacles. The manubrium has 4 simple lips. The gonads are developed upon the sides of the
manubrium and upon the 8 main stems of the radial-canals.

The type species is N. brachiatuiii Mayer, of the Bahamas and Tortugas, Florida. It
is the only species of the genus.

Netocertoides brachiatum Mayer.
Plate 27, figs. 4-6.

Nctocertoidei brachialum, MAYF.R, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 45, figs. 43-44, plate 18;
Memoirs Nat. Sci. Museum Brooklyn Institute Arts and Sci., vol. i, p. 12, fig. 7, plate I.

Bell 4 to 5 mm. high, miter-shaped, walls quite thin. 10 hollow tapering tentacles; one
at base of each of the 16 radial-canals. 1 6 to 25 small tentacles, one or two between each pan
of long ones. Longest tentacles about one-fourth as long as bell-height, their ends tightly

230 MEDUS.E OF THE WORLD.

coiled. The short tentacles are hardly more than cirri. No marginal sense-organs. Velum
well developed. 8 main radial-canals arise from the inanubnum, but each bifurcates, giving in
all 16 terminal-canals, which extend straight toward the circular vessel. The 8 proximal roots
of the radial-canals are bound to the manubriuin by groined arches. Manubrium broad and
disk-like. The mouth is at the extremity of a short neck, surrounded by 4 simple lips.
Gonads along the main radial-canals, adjacent to the stomach. Entoderm of tentacles,
radial-canals, and manubnum rosin-colored. Quite common at the Bahamas, but rare at the
Tortugas, Florida. Seen only in summer.

This medusa is closely related to Dichotomic cannoides, but in NctocertoiJfs 8 main
radial-canals arise from the stomach instead of 4 as in Dichotomic, and the manner ot branch-
ing is quite different, even in young medusae.

Monobrachium parasiticum Mereschkowsky.

Monabrachium parasiticum, MERESCHKOWSKV, 1877, Annals and Mag. Nat. Hist., ser. 4, vol. zo, p. 220, plates 5, 6. — WAGNER,
1889, Zool. Anzeiger, Jahrg. 12, p. 1 16; 1890, Archiv. Biologic, tome lo, p. 273, plates 8, 9 (details of histology).

This interesting hydroid has only I tentacle. Its medusiform gonophore belongs appar-
ently to the Thaumantiadae, but the free medusa is unknown, if, indeed, it be set free. The
medusiform gonophore is oval and is developed in a sac which arises from the hydrorhiza.
The hydroid grows upon the posterior part of the living shell of Tellina calcarea, near the
siphon. The hydrorhiza is a flat expansion of anastomosing tubes covered with a thin chitmous
membrane. The polypites are about 2 mm. long, tubular, with a single tentacle. This is 4 to
6 mm. long, tapering, lash-like, and arises from the side of the hydranth at a point about one-
third the length of the polypite from the mouth. The gonophores are medusiform, about as
long as the polypites, and they arise from the hydrorhiza by a short stalk. The contained
medusa is oval with a short conical manubriuin, 4 radial-canals, each with 2 gonads upon its
sides. Ring-canal and 16 rudimentary tentacles. Color of colony, yellow. White Sea, Russia.

K'l'I.X TU Vul.rMK* T AM) II.

Adiocheteumena, 430

•/•>'"" 15. 448, -/-/<;
Miulrs, 4^4
alternans, /-,'}
brunnea, ./f }
canarK-nsis, 452

' '""' ", 45°. 45'
1.1, hscholtzii, 450, ^•,",'
lacK-u, 453
pachyderma, 473
r/inJina, 450, ./,-_'

IOSI a, 451

.•Ks;im-lhi, 454

bitentaculata, 455

disMiiK-ma, 456
.K"Mi.-ra, 432, 467

o.rona, 437

H.n I-MVMS, 454

gemmifera, 480

glubosa, 476

paiip.-mil.i, 432, 437

rost-a, 480

EginiJ*, 428, 430, ±tf
I 'i'i>iiii/is, us, 16, 448, i\n

n: tinodiscus, ./.'iVi
/Eginodorus, 485
Eginopsis, 15, 44^, 450, 454,

hensenii, 455

liiur, ntn, 472,
mediterranea, 456
UK itcnsii, 472
/>.;,/; \ili-rmn, /~ ,'
Eginura, 11^44^^ l"7
gnmalJii, 4^'S, y-o

IHilStl, £/2

lanzfrottf, 468, ^dp
tn \iisnru, /f'*i
\\clu-ri, 470
Equorea, \i, ;j/

, ?;/

ciliata, 325
, .-;';, ,;. ; ;/
cubana, 325
cyanea, 326
iTamigramma, 445
discus, 325
i-iu-liroma, 222
fiiiliiulina, 326
flonjana, Jjo
forbcsian.n. ?2^
for sk/il, -a, ;_'•,"

Volume I extends from paj;r i to 2-0, volume Hi
Principal rcfcrt-iurv arc in italics.
/Equon

•17-

globularis,

"in nlamlira,

maldivensis, ,",'". ^'i

Illl S(.|H 111. I, ^^

mina, 12^

M "k .1, 32'', i-V

a, ,;,•/, \ppi'inli\, 41/7

, ,

pl< -1111111.11.1, ^X
puiu-tata, 444
pnrpuii-.i, ^S
i ii.»!ci|iinia, 4^2, 4^7

risso, 325

nvsoan.i, 32^
rtist .1, • • •
st.iui(ii;l\ pi) a, ^^
It-nuts, ;;_'

vitrina,

, i i. 196, , in
\L;.IS-.]/. \l. \.nulc-r, ack

i, I ), 2,'I, _','/

. alii nlata, 234

• ' ' /

I ••/.iitt/tii, 14, /(>/
conica, 402
tligitiilt1, /"-'

Hit, 11111-1/1:1, /"/

-t-in . '/, i itlt-ntnlis, ./a /
elongata, /"/

'•it! it;, rn, /<i /

i !;"<•", /"i
i, 402

.lflllll',1, 14, ,'ry-

hemistoma, ^j ,

/.//. I ».;, /OO

. /en
Ol hii'unn, fl'l

\" in. ill.ii.i, 2iS

W«, c^. w>

•;/n, 362*
\nllsi-ta, 4OI

I, 404

i !i>iii;ata, 404
405

[i]

INDEX.

Amalthtea, 5, 22

am&bigera, 22

junuarii, 2J

sarsii, 22

uvifera, 22

vardoensis, 23
Ametrangia, 224

hemisphierica, 224
Amphinema, 108

apicatum, 109, 112

nustralis, Appendix, 490,

dinema, 109

titania, 109

turrida, Appendix, 490
Amphocodon, 37

ampliipleurus, 39

fritillaria, 39

globosus, 39

gravidum, 39

unicus, 42
Amphogona, 14, 405

apsteini, 4.05

pusilla, Appendix, 497
Anthomeduste, 17

development, 3, 5
Atractylis, 201
Auliscus pulcher, 43

Batliycodon, 73

pyramis, 79
Berenice, 221

capillata, 222

euchroma, 228

huxleyi, 222
Berenix, 221

thalassina, 228
Blackjordia, II, 232, 2j6

manhattensis, 2JJ

virginica, 2ff
Blastothela, 29
Botrynema, 14, 394-

brucei, 395
Bougainvillea mediterranea, 133

paradoxica, 162

ratnosa, 161
Bougainvillia, 2, 3, 8, I$$

allmani, 160

nutiimnalis, l6g

bella, 161

benedenii, 161

bntannica, 161

carolmensis, 165

dinema, 164.

flaviJa, l6S

frondosa, 171

fruticosa, 160, Appendix, 493

fuha, 160, Appendix, 492

gibbsi, 169

glorietta, 172

kollikeri, 179

maclovtana, 160

Bougainvillia maniculata, ijo

multicilia, 164

muscus, 155, 161

niobe, 166

nordgaardu, 168

obscura, 172

octopunctata, 177

platygaster, 165

pnncipis, 160

prohjera, Ifo

pyramidata, 168

rugosa, If I

supercilians, 162

tnnema, IJI

xantha, 165
Budding in Medusa;: Hybocodon, 39; Sarsia, 48,

61-64; Margelopsis, 80; Lleutheria, 95;

Cytaeis, 133; Bougainvillia, 163, 166, 171;

Rathkea, 178, 179, 182, Appendix, 493; Pro-

boscydactyln, 191, 192; Phialidium mccradyi,

271; Eucheilota paradoxica, 285; Eirene

medusifera, 413; Gonionemus larvae, 345;

Limnocnida, 370; Narcomedusae, 429; Pe-

gantha, 447; Cunoctantha, 466; Cunina, 478,

481; Rathkea, Appendix, 493.
Bythotiara, 8, 185

murrayi, 185
Bythotiaridi, 183

Calcydion formosum, 136
Callitiara, 146

polyoptlialina, 147
Calycopsis, J, /jo, Appendix, 491

typa, 131 Appendix, 491
Calyptra, 376

umbilicata, 378
Campalecium, 291

meJusiferum, 291
Campanella, 207, 449, 454

pachyderma, 473

Campanopsis, 196, 295, 2gS, 302, ^OJ.
Campanularia bilabiata, 233

caliculata, 234

compressa, 233

dichotoma, 246

dumosa, 201

flabellata, 249

gelatinosa, 255

geniculata, 244, 246, 249

inconspicua, 266, 269

johnstoni, 262

noliformis, 265

synnga, 269

volubilis, 262
Campanulina, 234, 266, 314, 329

acuminata, 314, 316

hincksii, 284

languida, 269

repens, 235, 317
Cannota, 10, 197, 213, 221

dodicantha. 221

INDKX.

Cannotidae, 102, 196
Carmarina, 424

fungiformis, 425

hastata, 425

hexaphylla, 425
Carmaris, 424

giltschii, 425
lj 425

umbella, 425
( '.n mimothe beroe, 105
Catablema, I2O

campanula, 128

i un sronia, 128

vesicanum, 126
Charybdea bitentaculata, 455

ci<;«i<-ii«, 8, 182

centripetalis, 182
Chromatonema rubrum, i , ,
Cii , 401

camtschatica, 402

elongata, 404

inveitrns, 404

rosea, 402
Cir< ead e, 396
Circi-lla digitalis, 402
Cladocanna, 228

polyclada, 228

rhalassina, 228
Cladonema, 3, 7, 98

perkinsn, IOI
radiatum, <)<)
Cladonemida, 6, 83

i '.I, i.', it, ll,i, pj

prolifera, gq.
Clavttln, Appendix, 491

tontata, Appi-ndix, 491

gossei, 122

, ta, Appendix, 491

fi.if'ii.i. Appendix, 401
( '!,•, .dura tricuspidata, 122
r.l\ti,i, 10, 232, 261

bakori, 262, 495

bicophora, 262

compressa, 233

ri lindrica, 262, 265

eucophora, 236

flavidula, 266

falleata, 264

fragilis, 262

geniculata, 262

johnstoni, 262

polynesiz, 264

rriiit;irnir, 26$

simplex, 262

innvi-rsitatis, 265

i"ilnliilis, 262
Cotionidir, 5, '0
Codonium, 47

apicuhim, 59

conicum, 58

codonophorum, 61

gl niiiiili-rum, 62

I'linccps, 60
Codonorchis, 108
•• 'Irus, 1 1 1
Colobonema, 383

sericeuni, 384, 385

tvpicum, 385
('.'•nis, 7, /JO

iphthalma, 130

initrntii, /JO
t'onvt i{;i nrc of: C'tcnaria, t'ra.s|<i-dnt«-lla, and

Pelagothuria, J>4; Narcomedusac and Lepto-

lina nuilii--.!-, 15,84; Mydim rcn.i and C'tui»-
plidra-, 4<iO; Ma-risin and Hydra, 488
Corydendrium minor, 122
<moTpha, 22, j/, 37
annulicornis, 38
fririllaria, 38
januarii, 23
nana, 35
rmtiins, ?/, 41
ptnJuIa, 1 1

11\ il'iT.I, 22

vuL'iil.n.i, 35

C'orynr, 47

i-\iniia, 57

mirabilis, 53

prl.ii;ii-a, 89

producta, 63

rosaria, 59

Corynetcs agassizn, 72
Corynidae, 19
Corynttis, 6, 71

agassizn, J2

,n, tltfl, 7_J

coeruleus, 7J
Cosmetira, 201, 231, 261

^1', s, ll,i. : ;

snlinarum, 202
Craspcdacitsta, 13, J<5j

_
sewerbn, ?OT

Crnspedotit, 1 6, 1J
Crematostoma, 324

flava, 326
Crossota, 14, 595

brunnea, ^g6

norwegica,
Cn ptocarpx, 17
Ctenaria, 6, <)8

[>hora, <)8
Ciiliiin, 12, JJO

nphrndite, ^^1
Cubogastcr, 132

dissimcmn, 1^4

gemmascens, 133, 178

i-s, 449
primigcnia, 449

[iii]

INDEX.

Cunantlmhi-, 447
Cunarcha, 449

aeginoides, 454
Cuiiinn, 15, 448, 460, 473

albescens, 482

campanulata, 474- 47 ^

dodeeimlobata, 476

Juplicdta, 475, jSr

gl'jbosa, 474, 4~(>

kollikiri, 464

liitii'cntns, 474, 476

moncta, 482

mui ilaginosa, 475. -/Xi

multifJia, •!'•;
nana, 461

parasitica, 4(15

pi nvrinn, 4^ f

proboscidea, 474, (.76

prolifera, 474,

rliododactyla, 480

solinaris, 482

striata, 449

vitrea, 476
Cunissii, 1 6, 448, j.Sj

pnlvplii-rti, />'•,"

polyporpa, / ^ j
i.iiitliu, 15, 448, ./tfo

fowlen, 466

incisa, 472

uiirin, 46 r, 404
r.ir. koJllkeri, 464.

ftiirnsitifi, 46^

polygona, 465

tenella, 467
Cunoctona, 467

grimaldi var. munda, 470, 472

gumensis, 470, 472

lanzcrotae, 469

nausithoe, 469

obscura, 470, 472
CuspiJt'Un, 2111, 204, 286, J.S'.V
Cuvii-rifi, Id, 197, 221

cansochroma, 222

euchroma, 228

huxleyt, 222

Cyanea bougainvillii, 156, 160
Cytaeandra, 1^5

areolata, 140, 154

polystyla, 140
Cyt<ri.<, 7,131, 132, 17,-

exigua, 136

gracilis, 141

herdmani, /?,

macrogaster, 133

minima, 140

nigritina, I 5^

octopunctata, 177

polystyla, 143

pusilln, /?-/

tetrast\la, il]

vulgaris, 134, Appendix, 492

[iv]

Cytology: Amalthaea, 22; Pennaria, 26; Steen-
strupia, 31; Hybocodon, 41 ; Margelopsis, 81 ;
Eleutheria, 95; Cladonema, 100; Turntopsis,
144; Oceania, 148; GonionL-mus, ^48; Lir-
iope, 414; Geryonia, 427.

Degenerate medusae: Pachycordyle, 21 ; Bougain-
villia mamculata, 170; Eucopella, 233; Agas-
tra, 234; Eucopc parvigastra, 238; (lastrodrs,
461.

DenJroclava Johrnii, n~, 118, Appendix, 4110

Dendronema, 7, 102
stylodendron, 102

Dendrostaurinee, 8, 7^9

Development of : Millepora alcicornis, 16; IVnna-
ria tiarclla, 26, 27, Appendix, 487; Tridio-
rhiza, 29; Steenstrupia rubra, 32; Hybocodon
prolifer, 39; Sarsia, 52-64; Stauridiosarsia, '15;
Hydrichthys, 66; Ectopleura, 69; Connitis,
72; Margelopsis, 81; Pelagohydra, 83; Zan-
clea, 87-90; Eleutheria, 95; Mnestra, 97;
Cladonema, loo; Stomotoca, 1 12, 114; 1'an-
dea, 118; Turris, 123, 124; Cytaeis, 133;
Podocoryne, 137; Turritopsis, 144; Oceania,
148; Stylactis, 150; Bougainvillia, 160, 162,
163, 166, 168, 171; Nemopsis, 174; Ratlikea,
178; Niobia, 187; Proboscidactyla, 190, 192;
Willsia, 193, 194; Laodicea,204; Melicertum,
207; Dipleurosoma, 226; Eucopella, 23.5;
Agastra, 234; Eucope, 235; Obelia, 244 -'.f v
Tiaropsis, 258; Clytia, 263; Phialidium, 267,
271; Gastroblasta, 280; Eucheilota, 285;
Mitrocoma, 287; Stauropliora, 292; Kutinia,
297, 304; Phortis, 307, 309; Eirenc medusi-
fera, 314; Tima, 318; ^Equorea, 328; Gonio-
nenius, 344; Cubaia, 351; Olindias, 355, ?s8;
Craspedacusta, 364; Microhydra, 366; l.im-
nocnida, 370; Rhopalonema, 379; Homie-
onema, 383; Aglaura, 396, 398; Liriope, 410,
414,421; (iervoni.i, 424, pfi; Narcomedusse,
428; Solmaris, 434, 437; Pegantha, 447;
Cunoctantha, 464, 46^; Cunina, 47^, 477, 481 ;
Ratlikea, Appendix, 493; Mtrrisia, Appendix,
488

Dianaea, 116, 265
bairdii, 319
bitentactilata, 294
conica, 118
digitale, 402
exigua, 413
funeraria, 380
lucullana, 314
proboscidalis, 42^5
rotunda, 124

Dichotomia, 10, 197, 223
cannoides, 223, 230

DicoJonium, 5, 44
adriaticum, 4.7
rornuttim, 46
dinema, 47
•ftoridana, 46

INDEX.

Dicodontum jeffersoni, -jt>
Dicranocanna, 188

turcillata, 189

Digestion, Intracellular, in C'laspnl.u !•
Digitak- medusa, 402
Dimorphism in hvclroids: IV-jinana am! <

morpl.a, -!,'; Mnc'iniu and Stamiilia,

^4, ui); Glaums ;mi! Sarsia, 72; lioii^.nn-

villia am! NYmcipsis. 172; \j;astia, 1:4:

Obclia, 256
I )im ma, 44, 108

occllatum. 4''
Dimmatell.i, 108

c.avus.i, 109
Diocheteumena, 430
Dipetasus,

digomnius, }(u

Dipklllol), 2S'l

paivum, 2^.;

Difleurosoma, in, i .

1,1,1'iksii, 214,

'i, hracea, ?-«
pacific a, 22j

t \ptfiini, 22^
Diplnra, 37
Diplimlla clavata, 76
Dipurena, 7^

cati-nata, 77

cervicata, 76

oiim-a, 76

dolichogaster, 77

K-nilis, <.\i, 77

fragilis, 77

lialttrata, 75

. i, ; i)

picta, 77
strangulata, 76

a, 7, I Ij
saphenella, //f

turfiil'i, rif>, \ppt-tidix,
Distribution of: Trafhyiiu-ilus.-^,

nula, males am! ttmalrs, ,-71
\)\ sc.innota, 188

ilvsilipltura, 189
Dysmorphosa, 1^5

carnea, 136

cundiioila. i

tiubia, 141

t'lilgurans, I ^

gemmifcra, 192

-i.u-ih's 141

minima, 181

mimita, 140

ti inns, 141 , 142

Ectopl

Jiini'iitu-ri, (it), Appi'inlix, 4^'S
mintrva, 70

ochracea, 'HJ, Appendix, 4^X
pacitica, 70

i; I.ir

Eirene, II, 232, 2^, ;//
danduensis, ,'/ ;

naif, 402

gibbosa, 307

kambara, 309
medusifera, ;/ ?
fiellucttlii, Apprndi ...
guadrigatum,

J'l/IS, 7/J

riiiJiiln, ;//
Eireninte, i i , '
Eleuthena,

clap

Jni,

• allentini,

I miruiimi-iit, <Hnt of: light on regcm ration ol
I ml, . '7; Ilimn.' , upon IJou-

. 2 ;.(, < )l>
'•
cymbaloides,

loll. ,: -f«4

IIU'C I.lll'. 1, 27 I

rang

l-.ii, li.-il'itH, II, > ^2,
link, n, 2o2, .'//'/

bcrmudensis, .
JuoJecimalit,

111::, II '

/''"

. .-'$J

onium, - . <
68
!>f, 10, 231, • ; /. • ^>, 2'n

i(l2

nafa, 241)
annulate,
artii ulaia, 246
campanulata,
diaph

tllSS''*

folk/and

fll-il' . 2^2

••nn!!;ria, 2X5

f/'/

hyaltna,

>.|iln|ua, 250
parvigaslra,

pul\ '.So

polygena, 2f~, 2^1

tyla, 155
p' nt'oiniis, 247
••,//,/, 10, 2^1. j,-j
/i;/(."
campanularia, 2 ^

i n-nala, 2 ^4

. I'"1. l')'\
l-.iioipium. 2^4
parv iu.i-.ttum.

INDEX.

Eucopium pictum, 236

primordiale, 236
Kucoryne, 23

elegans, 25
Eudendrium, ettYct of light i.n regeneration ol,

ramosum, 161
Euphysa, 29
aurata, 35
globator, 24
mediti.Trani.-a, 35
u-ntaculata, 53
utrabrachia, 36
virgulata, 35
Euphysora, 29
bigelowi, 36
tetrabrachia, 36
Eurybiopsis anisostyla, 414
Eurystunia rubiginosum, 480
Eiitima, H, 232, 2i)$

brownei, Appendix, 496

campanulata, 29(1, ,'<>->, Appendix, 496

cuenilea, 296, 304

cuculata, 296, Ji/>

, nrva, 296, JOO

elephas, 296, joo

emarginata, 295

gentiana, 296, 502

gracilis, 295

insignis, 296, 299

lactea, 296, 500

levuka, 296, J07

<uar. occellata, 302
limpida, 295
mira, 295, 296
modesta, Appendix, 41)"
orientals, 296, 299
pretiosa, 296, jo6
pyramidalis, 308
variabilis, 312
I utimalphes, 295

brownei, Appendix, 41/1
coerulea, 304
indicans, 306
pretiosa, 305
scintillans, 306
Eutimeta, 295

gentiana, 302
lactea, 300
levuka, 301

Eutimina, 1 1, 232, Jol
Eutunium, II, 232, ?()$
elephas, 300
scintillans, 306
serpentinum, 300
soci alts, 506
Eutonina, 305
socialis, 306

Favonia, 153, 17-
bachei, 173

27

Fission or scissiparity in Obelia, 244, 249, 251;

in Gastroblasta raffa;lei, 280
Kovi-olia, 460

diadi-ma, 445

mollicina, 445

octonaria, 4<>i
l-'n-sli- water me

Thaumantias lacustris, [99;

, 363-365; Microhydra, 366;
Linincumla, 370; Moerisia, Appendix, 488
Fusion of blastuhr in Mitrocoma aniuc, 288

1,1, ii, 2}2, 2jg
ovali s, ?Sl

I ,://.!/, I. 280
tllllltla, 2f<)
( 1. 1 st null's, 460

paiasiticum, 461
ii.-mtnaria, 85
cladophora, 90
dichotonia, i)l
•-88

implexa, 89
nitida, 91
sagittaria, 88

( i. nil cells, origin of: IVnnaria, 25, 146; Dend-
roclava, 117,490; Podocoryne, 136, 139; Hou-
gainvillia, 162, 163; Obelia, 239, 251, 253,
254; Cunina, 479; Sarsia, Appendix, 488
Geryoncs, 424
(/. i v"'"'', 14, 4°0. •/-•/
appendiculata, 413
bicolor, 413
hiu-ntaculata, 294
dianasa, 425

rxigua, 413, 420

fungiformis, 425

hastata, 425

hexaph}'lla, 425

in. \icana, 425

octona, 123

pellucida, 314

probosciJalis, 4.25

rosacea, 417

tetraphylla, 417, 418
G, i yontJte, 14. -f°9
( ;, n.mopsis, 311

forbesii, 311

gentiana, 302

pellucida, 314

Glands in I.ymnorea alexandri, 75, 155
Globiceps, 23

globator, 24

tiarella, 25
Glossocodon, 410

catharinensis, 421

liajckt-lii, 421

liitkenii, 418
mucronatum, 413

tenuirostris, 416, 421
Glossoconus, 410

[vi]

[NDEX.

Glossoconus canariensis, 418

Gonads, position of, secondary : of Slabbcria lialttr-

at;i, 76; perradial, in Katlikca, i<So; IIU<M

in Obclia, 246, 253, 254; in ' . 103

Gonionema, 541

murbachii, 343

vertciis, habits ot, , / ,'
Goncomcandrus, 21 v

chiysostephanus, 220
( ionioiK-mouks, 350

U<"phil.i, ,o'
Gontonemus, I2,j^l

agassizu, 342.

J?{>r?sstis, 342, ,'/V

Kraftin}; of, j/i)

home Mi, 342, ;_y>

iniiilnii-liii, ^42, ,'/ ,\ 41 (S
ins svstt-ni, .|4''

pilagicus, 342, 349
a nsis, 342, ;/i;

U2, ?«

( lon\ nenia, 341

' .iroa, 293

mirabilis, 294
Gossea, 13, jd/

brack \'in

circiniata, 367
lu-tt-s, 367
(iHmtli, r:ite of in hydroids: Sarsia, 5^; I5ou-

gamvilha, 162; Obeli. i, 246
( lymnopthalmz, 17

Habits: Collapsing of Dipleurosoma,

la\ing of Stomotcca, Il.j; ( )i-r:.ni.i, 14^;

.Equorea, 328; < lonioMcmus, ,'^4; l.iriopc,

414; Geryonia, 42^, I-'i-nlui!; of /.uu Ic .1. 89

Turns, 126; Swimming of Obelia, 24^; (ion-

ionemus, 344; Cubaia, 351; Olindias, 355;

Craspedacusta, 365
Halatractus, 31
llnlcordyle tiarella, 25
llalicalyx, 352

tenuis, 354
Htiltfreas, 14, }<?0

alba, 393

coined, jgj.

glabrum, Jp2

minimum, 391

ftapillcisutn, 391

rncovttzie, jpj

rotunJatum, 3Q2
llaliscera, 390

conica, 394
Halitiara, 105

formosa, 107
IJalitrephfS, 59^

in u as i, JQ./
Halmoniisis, 198, 199

lacustris, 199
Halocharis, 85

spiralis, 72

/'sis, 286, j_'j

.inn.i-, 287

cruciata, 277, 285, 287,

megalotis, 289

ocellata, 325
I -tbella calcarata, 202
Hermaphroditism: I-U utlu-na. 84; i

99; Amphogona, 405, Appendix,
Heterostephanus, 29
Hiterotiara, 7, 707

anrjnymu, 707, Ap|H-iuli\.
Ilippocrene, 155

aurca, 162, 164

bougainvillii, .

biitannu .

carolim IIM\, i'i^

criu >

ma. 160

I'liiKtata,

platynastt r, 105

p\ raniid.ita, i

Mlpt'li lll.lils. l62

. '4. 377,

•ill'*', 393
amflum,

rlon^atum, 379
ni:i. rogasti i ,
null
fl. :

it/a-, ^(^
tyfifitin,

ampliipli in US, }Q

.iilii-onn-;, 39
, liil.mif, fj
: linstni.i-. /_'
forbesii, /_•

n

.Inns, 39
.

oividi-ntalis, 42

//

, 41
firnlifi i ,
ftii

II 111: U

1 1\ ilraitc n.i, 4^S
Ilyilrai-tinia au.ilaia, 140

nun

> ,15, 44^, /.is'

salenskii,

In in •, .;i i

[vii]

i lea, 304
ijibbosa, 507
mollis, ^u
palki-nsis, 309
pellucida, 314, Appendix, 496

INDEX.

Irene viridula, 31 I
lieninpsis primnidialis, 351
Irenium, .; i i

quadrigatum, 313

/1,-ti'if^iis, 1 I, 232, ?/('

/(, \nn, mahs, ;/<>
Isonema, 3X3,

amplum,

Kermin IKTIH , I 22
Kollikilia fascicuh'ta, 171)
Lat'ci-a, 201

caK.nala, 204

coinuta, 2OI
Laodice, 2OI

eellulaiia, iijli, I'ji)

indi. a, 2 i2, i"4
Laodii . .'•-,, -'"i

caK-aiat.i, J i, ' '4

( i ! rhapinam, ' 6

CTU ata, 'Ol

( ' I • Vk li|)l , 2O6

i i i !> mlis, 206
fijiana, .?<>•,"

vai. indica, 205
maasi, 205
mai.i;n:\ '. '. 204
( ?) nepluna, 206
piii, hra, 202, 2oj
ulothrix, 202, 204
Laoniedi a, i^N

dichotonia, 244, 24^, 255

divaricata, ' i ;

m l.innosa. 244, 247

•M 1M. 111. II. I, 24U

Lar sabellarum, 193
Leptomt-ditsie, g, /i/>, 231
Lessonia, 397
Leuckartia, 424

proboscidalis, 425
,13, 370

: iml :Li', 13, l6g
Limnocodium, 303
sow riliii, 3')^
victoria, <'••;
Limnorea, i ^ ,'

IKII \cgica, 154
proboscidea, i ^ ^
Liriantha, 410

catharinensis, 421
nuR-ion.it::. (14
n iiaplu-lla. 418
Linof>, , 14, 4.09, //"

appcndiculata, 411, 413

. 41 1, jiS

catharinensis, 412, 421
rasiformis, 411, -/r?
cerasus. ;i <
omipacta, 417

s, 412, jit)

crucifera, 417
ilistiinognna, 412, 420
,'iu \lnii, 412, /_'«
.•\igiHi, 411, ^/J
li,i,;-ki-lti, 412, _/J/, Appi-ndi\,
li\ralina, 421
/; yperboltca, -j i<<
unli, ,1, 41 2, ^-?;
lifiutina, 420
lntk,-nii, 412, //.V
minium, 412, 7-'^

»/ll, inimtil, 411, 413, /'-/

proboscidalis, 425

ii /''I, ^f-, AppL'iulix, 4117
r, utigera, 412, 413, 411;, /-'/
tenuirostris, 41(1

ti'<rn[>li\ll<i, 41 2, ]lS

Liriopsis campanulata, 302
Li/iisa, 155

multicilia, 1^14

octocilia, 161

prolitVra, 170
Lizzclla, 1/5

octdla, 180
I.i//.ia, 175

blondina, 181

clapau-dci, iSi, Appendix, 4(^

formosissima, 177

lirata, 177, 17.1

kiillikeri. 171)

octopunctata, 177
Lymnorea, X, /,"j

all \iinji', ijj, Appendix, 41/2

hori-iil'i, 140, /_,/

dibaha, i^

tiiiii-r/istninii, 7 _•>•_,

_
rrmlra, 153

Mahella gracilis. i JQ
\tisotias, 13, ^i^X

Margelidic, 102

Mill i'f/n;,r, / ,;

\ [argelis, i ^

autumnalis, 169
carolmensis, 165, 167
nordgaardii, 168
principis, 160
rannisa, 161
trinema, 171
zygonema, 164

Margellium, 175
grata, 179
octopunctatum, 177

'Mll,r. 6, 2O

ris, fi, ?0

[viii]

hteckelit, So
hartlaubn, 82
stylostoma, Si

IXDKX.

Marmanema, 376
clavigerum, 382
mammseforme, ^

n inpaniun, ,
MH -radia, I y$, '54
Mi-dusa xquorca, 202, 324,

bimorpha, 522

campanulata, > iX

l-n-llllll pl'lisilr. '.{ ^

cmciata, 201, 202
cymbaloidea, 2'>X
i;li>liul.nis. ^^
hemisphaerica,
iiuinna, 1 .[X
mucilaginosa, 4X1
patina, ,'25
pilrata, 123
proboscidalis, 41^
saltatrix, 210, 22O

\lrliivtta, 2O/

digitale, 402
fasciculata, 17*)

plrmostoin.i, 146

Melicertella panocto, 209, 210
Melicertidium, 207, 209

campanula, 207

octocostatum, 208

\l,h, iitnite, 9, IQJ
M.li.-crtissn, (), 197, 2O<)
fltii'tgi rn, Jlo

malaytca, 211

Mi-li, ,itnm, 9, 197, so'/, 2IO

fiinipnmi/ii, ji)~

p, orgii '-mi, soy

nctncostatiin. .

pnnocto, 209

penicillatum, 218

( ':) proboscifcr, 209
Mi-siun in.i, 211, ^24

abbrt-viata, ^^

li.iiiilu, 326

ccerulescens, 326

tlubium, 32^;
eurystoma, 325, 327

nUKTOll.il t\ 1.1, H ,'

pi-ncile, 333
|)ilcus, 211
victoria, 330

.nnfinnii, ^. /{

' I I

Mi: rohydra, i i,, ;iV>

n, ^66
MiiTostoma, 92

ambigua, 92
Milli-fora, 16

',rnis, 16

Mill, porinte, mrdus.r <>f, ir>
n, i r, 232, 25>'.

unn.r, 287, 288, 290

Mltt'i. ',111:1 : III, It.!, .

cruct.ita,
discotdea,
lenJenjeldi, 290

i/uta, 289
mtnerva,
polydiademata,

Mill-in diiiclla, 2S«i

fulva, 290

pi)l\ 290

Minocomiiiin, jX'i

cirratuin, iSS

parasites,

'i-iia, 71, 14^, 14(1, 14-
isin, Appnulix, 4SX
l\'ntisi. Appendix, 4X8

Malli, 111,1 nil diisa. 445

a< liiiiin j

Monocaulus prinlulus, 41
Multioralis,

Cl\ .|]|S, 2Xl

'iii;i[i of, }, i ^.
/"", v X, Xi.i, ;-_•, \ppiiuli\,

'"•< I, '7i
f I Hi I j, 111. lf^

"ililn-sii, 82, 173
liiti-rnni ma, I 7^

tt>,:</i-s, to, 197.
brachiatum, 221)
Nii;ritina, I 52
;,\-

dendrotentacula, i ^~
Nucli-itVi.i-, 103

Oln-liii, 4. io, 2^.

[ix]

ii>ii;i, ' .
mill: nils, 2<j6
nustralis, 24'', J./-
nii.stri>-»i-iu^i:i-, 254, 255
bii-uspidata, 254
bidentata,

' -,4. Appt-ndix, /,;,'
bit'un-ata. 4115

ill', - /it
chinentis,
commissuralit, J//, Appendix.

, until,.,

i

diapliana, 24^, 2^2
dit'li'il'iniii, 244, j^rt
ill\ aiu-af :i, -147
Juliiii, 24'\
H.ihi-llata, 241;
fnigilis, JiJ
gelatinosa, _•//

INDEX.

Obeli a gracihs, 253

griffini, 252

gymnophthalma, 244

helgolandica, 253

hyalina, 248

leucostyla, 244

linearis, 257

longissima, 255

luciiera, 249

nigra, 256

nigrocaulus, Appendix, jyj.

plana, 2./p

filicata, 2$6

polystyla, 255

purpurea, 243, 256, Appendix, 494

pyriformis, 24.7

rhunicola, 246

serratula, 257

sphrcrulina, 246

striata, 24.8

surculans, 2$2
Ol.Jitur, 10, 231, 232
Oceania, 8, I2O, 146, 265

ambigua, 274

armata, IJ-J

blumenbachii, 177

brunescens, 274

carolinae, 275

coccinia, 123

conica, 118

coronata, 123

dinema, 109

discoidea, 272

flavidula, 147, 266

folleata, 264

gelatinosa, 272

globosa, 122, 272

languida, 269

magnifica, 269

mccradyi, 271

octocostata, 208

octona, 123

pacifica, 273

phosphorica, 355

pileata, 123

polycirrha, 143

saltatona, Il8

sedicimcostata, 1 18

singularis, 273

tetranema, 106

tubulosa, 52

turrita, 123

viridula, 311
Oceanidts, 7, 17, 102
Oceanopsis, 281

bermudensis, 282
Octocanna, 12, 320

octonema, 325

polynema, 320
Octogonade, 12, J2I

mediterranea, 322

Octonema, 2OI

eucope, 206

gelatinosa, 214
Octorchandra, 295

canariensis, 302

germanica, 302

orientalis, 496

vanabihs, 312
Octorchidium, 305

tetranema, 302, 306
Octorchis, 295

campanulatus, 302

gegenbauri, 302

orientalis, 299
Octorphalon, 2OI

fertilis, 206
Oli n Jiada, 12, 340
Olindias, 12, JJ2

malayensis, 353, 356

mulleri, 355

phosphorica, 353, 555

saml/i/ijuiensis, 353, y^

singularis, 353, 357

tennis, 353, 354
Olindioides, 13, 358

formosa, 358
Orchistoma, 9, 197, 211

agartcifortne, 212

pileus, 211

steenstrupif, 21 1

tentaculata, 212

Pachycordylf, 5, 20, 21

Jcgeneratus, 21

weistnanni, 21
Pachysoma flavescens, 434
Pandasa, 1 16
Pandea, J, 116

conica, 118

flavidula, 147

maasi, III)

minima, 118

saltatona, Il8

violacea, ug, Appendix, 490
Pantachogon, 14, 388

apsteini, 405

hcsckelii, 381)

rubrum, 381)
Parasitism: Hydrichtliys, 19, 6j; Mnestra, 96;

Stylactis minoi, 149; Cunina, 379, 425, 427,

429; Cunoctantha, 461, 4.64., 4.65; Gastrodes,

461

Parvanemus, 21
Pectanthis, 372

asteroides, 374
Pectis, 372

antarctica, 375
Pectyllidae, 371
Pectyllis, 372

arctica, 372
Pegantha, 15, 430, 439

INDEX.

Pegantha aureola, 440

clara, 445

cyanostylis, 441, .;/,-

dactyletra, 440, / / /

dodecagon, i, y./ /

goJeffroyi, U>

Ln-is, 444

I unul at a, //<<

intigiti/ii it, ,j /<i

martagon, 44. ., //;

pantheon, 440, .//_•

punctata, 440, .///

quadriloba, 445

subuliln, {if

simplex, 44^

smaragdina, 441, •//(">

Iriloba, 440, .^/j

'•fn, ././/
l'i M mthid.e, 420, 4^1

Pi • .isia, 439

dodecagons, 444

Pelagic hydroids: Margelopsis, So; Pelagolmlra,

83

/'. lagohydra, 6, 83
mtrabilts, 83

.mi, 5, 23
adamsia, 2j
cavolini, 24

disticha, 2J, Appendix, 487
gibbosa, 25
mornata, 4^-
marina, 24
pacifica, 28
rosea, 2J

symmetrica, 24, 25
titir,lln, 2$, Appendix, 487
vitrea, 28, Appendix, 487
Perigommus, IOQ
cidaritis, IIJ
joncsn,'! 14
minutus, 124
napnlitanus, 109
ii pens, 124

schneideri, 109
steinachi, 109
sulphureus, 80
vagans, Appendix, 490

1','t wj, 14, 406

dissogonima, 407
incolorata,

luierna, 407
Petachnum, 360

tiaropsis, 361
I'et.isata, 360

eucope, 361

rabbeana, 361
Petasina:, 13, jdo
Petasus, 13, 360

atavus, 361

t, tfiinema, 361
I'liialella, 234

Pliialella t'alklandica, 237
I'/iialiJium, II, 232, 265

atnbiguum, 274.

ampullaceum, 123

brunescens, 2J^

buskianum, 2JO

cyrnbaloideum, 235

duodecimale, 2Kj

gelatmosum, 272

globosum, 2"J2

gregarium, 2J2

lieriiispha;ricum, 266

iridesccns, 273

languidum, 261), Appendix,

IIIIIKL-, Appendix, 495

pacificum, 273

simplex, :j^
singularis, 2J$
ic mporarium, 266

ii'iiue, 310

variabile, 235, 266, 280

variabilis, 270

PliiiiliniF, 232, 261
Phialis, 286

cruciata, 288
Phialium, 281

bakeri, Appendix, 495

dodccasema, 283

duodecimalis, 283
Phialopiis, 232, 266, 276, Appendix, 495

comata, 2j6

diegensis, Appendix, ./<y
I'liiiiliiciuin, II, 232, 2J^

Carolina, 275

comata, 276

mbengha, 275, 276

•clrens, 2/5, 276
Phorcynia, 324
Phortis, II, 232, 305,507

ceylonensis, jop

clliceana, jop

gibbosa, 307

kambara, jop

lactea, 308

fulkensii,

pyramidalis, 308
Phyllirhoe, 97
Plancia, 293

gracilis, 21/4

Platypyxia cylindrica, 265
Plotocnide, 51, 66, 71

burealis, 106

inccrta, 106
Podocoryne, 7, 7?5

ancchinata, Appendix, 492

areolata, 140

cornea, 136, Appendix, 492

conchicola, 136

delimit!, ;yj
Jit In' a, 1. 1 1

313

[Xi]

INDEX.

oi V.''1 falgurans, /j
gracilis, 141
humilis, ij.1, //J

nit n tit n, I^o
octostyla, I-/.O
f'j/VStvliI, I/O

t, inns, rji
Pnlvcanna, .524, 315

aim nc-ana, 32(1

crassa. 33(1

f'orskalea, 32(1

fungia,325

<;i •iiii.mica, 325

grcenlandica, ^-,

italica, 325 327

purpiirostoina. ^ .;

vitrina, 325
Polycolpa, 431

Imskalii, 433, 437

zonaria, 433

/.oiuiH-his, 433
J'iJ\'i», Ins, 9, 197, J/,S'

campanulata, 218

ininiitn, J [ty
pt'nii illiitit, 2lS
pinnatus, 218
saltatrix, 220

Polyxenia, 337, 439
albescens, 482
cvanogramma, 44,'
cyanolina, 445
cyanostylis, 445
flavesctns, 434
[eucostyla, 4^
pleuronota, 338

Pottsia, 366
ryderi, 366

1'rnl'tiS: liln, t\l,i, S, ;,\

brevicirrata, 189
flavtctrrata, iS()

var. stolonitVra,

occidental™,

var. gemmifera, i<)2
var. stolomfera, i<>i
tropica, 191
vanans, 194
Protiara, 7, 71. JOJ
beroe, 106
borealis, 106
formosa, io"j
ha'ckclii, 106

}'s,n,lofl\tia, II, 232, 2j8
gardineri, 2JQ
pentata, 265, 2-jS
mil, 6, f)2

ambiguum, 92
darwinii, Q2
Ptychogastria, 13, 57^

Ptychogastria asteroidt-s, 374

polnns, _^J
Ptychogastridee, 13, 577

Ptychogfna, 9, 197, Jl-/.

californica, Appendix, ./(>/
ervthroponon, J/7

/ o /

1,1, It'll, _'/ f

longtgona, 2ij ,

pinnulata, 215

var. intermedia, 215
Pulsation of: Comic pusilla, 51; Sarsia, 52;

Polyorcliis, 219; Obelia. 245; Craspedat-ust.i,

365; Gonionemus, 348; ( leryonia, 426
Purena, ~ ^

brownei. Appendix, 489

"emniifera, 62

strangulata, 76, 79

Rutliki-n, 8, 775

blumenbachiana, 177
blumenbachn, i~~

,!,•« uns, /iS'/

fnsi ii ulata, if(>

formosissima, ijf

octonemahs, iSn

octopuuitiitti, 177, Appendix, 495

var. grata, 179
Reactions to: light by Eudendrium, 4; Sarsia, 52;

mechanical stimuli by Rathkea, 180; ehemi-

cals and electric current by Polyorcliis, 219;

chemicals by Obelia, 243; mechanical stimuli

by C'osmetira, 261; photic and mechanical

stimuli by Eutimium, 306; light and other

stimuli by Gonionemus, 347; various stimuli

by Craspedacusta, 365; chemicals by Gery-

onia, 424, 427
Regeneration: Eudendrium, 4, 27; Pennaria, 25;

Obelia, 239; Tubularia, 239; Tima, 318;

Gonionemus, 346; Liriope, 410; Gcryonia,

427; Solmaris, 436
Rhacostomn, 524, 335

atlanticum. 335

dispar, 326
Rhegmatodes, 324

floridanus, 330

globosa, 332

lacteus, 333

tennis, 332
Rhopalonema, 14, J7<5

cltruigerum, 382
,ir uleum, v^'n

polydactylum, 379

striiituin, jSl

tvpicum, 578

1'i-lntiim, 377, 378
Rhopalonemtnee, 14, ^J-6

Sabellana, 112
Saphenella, 234

dissonema,

iri, II, 108, 232,

[xii]

INDEX.

Saplicnia balearica, 293

bitentaculata, -•<;/

dinema, ion, ii > /

gracilis, zyj

titania, 109
Snrsia, 5, ft

uiigtilata, 60

apicula, 59

ban.-nt.sii, 55

brachygaster, JQ

i-lavata, 77

cocci 'h, tra, App> mli\, 4XS

codonophora, 61
ont

ter, 77
, 57

flammea, 6f, 120

Xi'ininiffra, 6?

"ladalis, s ,'

gracilis, 60

hargitti, 63

macrorhyncha, 52

nil nun n, 59

mirabilis, 53

var. reticulata, 57

nodosa, 48

prince ps, 60

prohfera, 6l

f'lil.-lhlLi, 57, Appendix, 4S'X

radiata, vS1

resplendent, Appendix, 4X7

rosarta, 59

siphonophora, f>2

strangulata, 79

tubulosa, 52

turricula, 57

(Stauridiosarsia) prnJnt /
Sarsiantf, f, 20
Sarsiella, 44
Schizocladium, 2^S
Scrippsia, 197, \j>prn.li\, 4,^

[>,!,- lp, 'I, A|)|)ciuii\, 4., ^

Si -\ plus mucilaginosa, 4X1

St.ismial dimorphism: I'rnnaria. -!'', Saisia, 5^, 55

Sertularia dichotmna, 246

tulatinosa, 244

gcniculata, 249

longissima, 255

pennana, 24

volubilis, 262
Sexual dichromatism: Stomotoca dinema, 111;

< )n-histoma pilcus, 21 1
Sibogita, 8, 186

rtrica, 1 86

nauarchus, Vppendix, 4<)i

simulans, 187

/V/M, .\|i|)cndix, 491
s, 293

bitentaculatus, 294

insignis, 299
Slabbena, 6, 75

Slabberia brown, i, Appendix,

""'•', 77
halterata, 75
ophiogastt
pyramis,
strangulata, ~6

' . fragilis, "

Slllllltll,',!, 14,

leptogaster, ^-

•iipanuin, ^78

I ,. , • , ; ,'_>

. 444
corrj,,,,, 432, ;
coronantlia, 437
/?«',..,,;;>, 4^2, 433. /,'/, 4X4
fnrskulii. 432, ,/;;

•j. nhaiiri, 437
iiu-isa, 4 Si ^
nisflllpt.l, 4 tj
/.•»//, H/I;, 4^2, / ;V

ostyla, 4.32,
nuu-ila^iiiiisa, 4X1
iiniltil'Jiiitii, 4^a, f
punttara, 444, 4<S'4
Milinaiis, 4^.?. / ,--
Sul missus, 1 6, 44X, ,

III/': Si : II f, /\'j

/•/,', kii, '
ephesius,

j,il ;'ii, :

III: /.,,', fS ,'

in, it -//,;///, /A'/
Siilnioiu l:i, 4^1)

flavesi-ens, 4^4
S'ilnnin,!,-lln, 1^, 4(.S', /,-/

bitentaculata, /i,

hensenii, 455

»/, diterrdnt :i, 4-> v /i"''
Solmundus,

elacialis, 472

tetralinns, 4^2

retrain mils, 472
SpirocoJon, 9, 197,

sultatrix, 22O
Stauraglaura, 14, /cA

1,'tl ,li"ll!: IIK1, jOll

Stauridia producta, 65

Stauiiilii-. 99
Si.iuijiliiini, 64,

productun
StauriJiosai '/•;. s. ('I

Stilur'iJis.lif. 0, !<)/, -1/?, 221, 22S
hit . f{

niprii-anv, 2*21
tftriisttinriis, 21^.

Staurophora, it, 2^2, 291

jillL-liltl,!:

kiitliii, •

laciniata,

fxiii]

INDKX.

Staurophora niertensn, 291
Staurostoma, 291

arctica, 292
Stft-nstrupta, 5, 2$

aitrata, 35

austrahs, J7

bigelowi, 36

cranoides, 31

galanthus, 31

globosa, 36

gracilis, 31

lineata, 31

occidentalis, 42

rubra, J7

tetrabrachia, 36

virgulatti, 55
Stenyo, 47
Stomobrachium, jJJ

lenticulare, $22

octocostatum, 208

stauroglyphum, 323

tentaculatum, 322
Stoinotoca, J, loS

apicata, 109, 112

atra, III

australis, 1 1 1

dinema, 109

divisa, 114., Appendix, 490

jonesii, ir4

octicJra, III, Appendix, 490

periphylla, 113

pterophylla, 77J

rugosa, 112
Stvliictis, 8, 149

arge, 757

fnriola, 141.)

hooperi, 750

minoi, 149

Syncoryna cleodora;, 122
Syncoryne densa, 53

eximia, 57

gravata, 52

linvtllei, 7'> 72

producta, 63

prolifera, 61

reticulata, 57

rosana, 5p

sarsii, 52, 60, Appendix, 488
Syndictyon, 47

angulatum, 53, 59, 60

reticulatum, 57

Tentacles, hydra-like tentacles of Moensia,

pendix, 488
Tetracannota, 224

collapsa, 226
Tetranema jeronauticum, 235

eucopium, 199
Tetrorchis, 388

erythrogaster, 388
Thalassanthea;, 428

Thamnitis, 8, 7J2

tctrella, I°j2
Thamnostoma, 153

macrostoma, 153
Thamnostylus, 8, IJI

dincma, 7J7
Thaumantiada, 9, /p6
T li.intnimti/is, 9, 197,

buskeana, 270

cellnlaria, 198, 7p£>

corynetes, 367

cymbaloides, 235, 311

eschschnltzti, 198

forbestt, IQQ

globosa, 235

hemispliaerica, 266

inconspicua, 266

lacustris, 7pp

leucostyla, 266

lucifera (lucida), 250

m&otica, 2OO

mediterranea, 2O2

melanops, 266

milleri, 208

multicirrata, 259

pilosella, 261

plana, 249

punctata, 355

quadrata, 200, 235

rubrum, igg

thompsoni, 199, 270

undulata, 202
Tholus eurygaster, 383
Tiara, 120

coniferi, 128

intermedia, 123, 125

oceanica, 125

octona, 123

papilis, 123

|iapua, 125, Appendix, 491

prismatica, 125

reticulata, 125

rotunda, 124

sarsii, 123

smaragdina, 123

superba, 126

Tiarella parthenopaea, 127, 147
Tiaricodon, 71

coeruleus, 73

Tiaridse, IO2

Tiarin<z, 103

Tiarops, 286

Ap- Tiaropsidium, 495

kelseyi, Appendix, 495
Tiar.psis, IO, 231, 257

Javisii, 259

diademata, 258, 260

indicans, 306

kelseyi, Appendix, 495

macleayi, 2$()

mediterranea, 260

[xiv]

201, 238, 259, 265

INDEX.

Tiaropsis multicirrata, 2$g

oligoplocama, 259

polydiademata, 290

punctata, 260

rosea, 260

scotia, 259
Tim,,, II, 232, 311,5/4

/M;I ,///', 315, 3/9

:l,,bris, 315, 3/9, Appendix, 497

formosa, 315, 5/7

lucidlana, 3/4, 315, Appendix, 497

prllucida, 311, 314

teuschen, 315, J/$>
Timoides, 9, 197, 2/2

agassizii, 212
Toxorchis, IO, 197, 22<?

arcuatus, 228

kellneri, 229

thalassitia, 228

Trachomedusae, 339
Traehymedusiz, 12, 559

origin of, 3, 339
Trachynema, 376, 382, 383, 401

camtschatica, 402

ciliatum, 378

digitalis, 402

eurygaster, 383

longiventris, 385

octonarium, 378
Trachynemidte, 13, 375
Trickorhixa, 5, 28

brunea, 21)
Tubularia dumortieri, 69, 70

prolifera, 43
Tubulurinc, 17
Tiirris, 7, 720

"Turris"=C/ovu/a, Appendix, 491

borealis, 402

brevicornif, 127
ampanula, 128

cceca, I2J

coniftra, I2S

digitalis, I2g

episcopalis, 126

etirystoma, 128

fontata, Appendix, 491

neghcta, J22

papua, 125

pelagica, I2f

pileata, 123

pnsmatica, 125

reticulata, 125

rotunda, 124

superba, 126

vesicaria, 126
Turritopsis, 8, 135, 1$$

lata, 146

nulricula, 136, 743

polycirrha, 143

Turritopsis polynema, 143

I'.illftitinia, 12, 352
falklanJica, 352

Variation and Aberration in: IVnnaha, 27;
Sarsia, 55, 57; Bougainvillia, 166; Nemopsis,
174; Ratlikt-a, iS2; Milucrtum, 2oS; Ui-
plciinisonia, 224; Obelia, 241; (_'l)tia, 265;
lidium, 270; 1'seudoL-luia, 27. j, ( )hndias,
^S; ln-iin|)sis, 311; .'l,i|ii- . 332;

Goniont-mus, 346; Olindias, ^S; 1 unnc.i-
nida, 371; Aglaura, 399; Liriope, 412; Sol-
niaris, 454; C'unina, 4X0; MHTIM;I, Appen-
dix, 489

Veiled Medusa:, 17

\V,ll,tta, 188

ornata, 1 89
Willia, 188

furcata, 193, 194

geninnlera, 192

01 < nil ntalis, 193
ornata, 189, 192

inn,,,di, is/

ff ill si, i, 9, 184, 1 88, /9J

brooksii, 195, ig6

mutabilis, IQ.J.

ornata, 189

st,-llata, /93

variuns, IQJ
Wrightia, 269

X.intlica M-utigi-ra, 421
ti-frapliylla, 4lS

7. ancle it, (>,

ainbigun, 92
cladophora, go
costata, 8j

Appendix, 4X1

inif'l \,,. .Vi;

nit id a, gi

L'opsis, 6, gi

dichotoma, gi
Zygocanna, 12, 337

costata, 338

pleuronota, 33$

purpurea,
Zygocannota, 337

purpurea, 338
Zygocarinttln, 12.

diploconus, 359

undulosa, 3^9

Zygodaetyla, 12, 324. 533
crassa, ^6
cubana, 325
cyanea, 326
granlandicus, ;;-,-
rosea, 325
vitrina, 32^

[XV]